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Full text of "Systematics of the Bufo coccifer complex (Anura: Bufonidae) of Mesoamerica"

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.Em 



Scientific Papers 



Natural History Museum 
The University of Kansas 



31 March 2005 N u mber 38: 1 -27 

Systematics of the Bufo coccifer Complex (Anura: Bufonidae) of 

Mesoamerica 



By 

Joseph R. Mendelson IIP, Beck^ L. Williams-, CnKisTorHEK A. Sheil^ 

AND Daniel G. Mulcahv* 



Ubi 



' Dcpaitiuoit of Hci-pctologi/, Zoo Atlanta, S(HI Cherokee Ave SE, Atlanta, GA 3U315 \'' •*^^-dc\TY 
-Departiiieut of Integrative Biologi/. University of California, Berkeley, C A 94720-iUO . .;;Vt-R^ 
^Departine)it of Biology, jolni Carroll University, University Heights, OH 44118 
^Department of Biology, Utah State University, Logan^lT 84322-5305 



CONTENTS 

ABSTRACT 2 

RESUMEN 2 

INTRODUCTION 2 

Acknowledgments 5 

MATERIALS AND METHODS 5 

Morphological Data 5 

Molecular Data 6 

ACCOUNTS OF SPECIES 6 

Biifo coccifer 6 

Bufo cycladen 11 

Bufo ibarrai 13 

Bufo pisinnus 15 

Bufo portcri 17 

Bufo signifer 20 

MORPHOMETRIC ANALYSES 22 

MOLECULAR ANALYSES 23 

LITERATURE CITED 24 

APPENDIX 26 



© Natural History Museum, Tlu' University of Kansas ElTlSt MfiVf Lf'TarV '^^^ ^'" "'''■*'"^*^2 

Museum of Compara*ve Zoology 
Harvard Unlversitv' 






] ds>^- 



Scientific Papers 



Natural History Museum 
The University of Kansas 



31 Mcirch 2003 Number 38:1-27 

Systematics of the Bufo coccifer Complex (Anura: Bufonidae) of 

Mesoamerica 

ubrary 

Joseph R. Mendelson III', Bhck^ L. Williams-, Christoi'hek A. Sheil\ 



ANL^ Daniel G. Mulcahv* 



- ^m 



' Dcimiiiiieiit of Ucrpctolo^ii, Zoo Atlniitu, S(H) Cherokee Ave SE, Atlanta, GA j03]5 H '-'•^y^-rrY 
-Department ofhite^rative Biology, University of California, Berkeley, CA 94720-3140 V. ''c-^^^ 
^Department of Biolo<;y, John Carroll Universiti/, University Hei;^hti, OH 4411S 
^Department of Biology, Utah State Universiti/, Logan, UT S4322-5305 



CONTENTS 

ABSTRACT 2 

RESUMEN 2 

INTRODUCTION 2 

Al K\'OVVLEDGMENTS 5 

MATERIALS AND METHODS 5 

Morphological Da la 5 

Molfxular Data 6 

ACCOUNTS OF SPECIES 6 

Biifo coccifer 6 

Bufo cycladen 11 

Bufo ibarrai 13 

Bufo pisiinius 15 

Bufo porteri 17 

Bufo signifcr 20 

MORPHOMETRIC ANALYSES 22 

MOLECULAR ANALYSES 23 

LITERATURE CITED 24 

APPENDIX 26 



© Natiiivil History Museum, The University of Kjnsds EmS^ MtiVf LF'TarV '^^^ ^'^' ' "^'■*""''*^2 

Museum of Compara*ye Zoo'o^ 
Harvard Univ6rs[t>' 



2 Scientific Papers, Naturai, Hisiorv Mi si,iiM,Tiii' Uni\i ksnv ui- Kansas 

ABSTRACT Many populations of toads occurrinj^ between west-central Mexico and Panama have been 
referred to Biifo coccifcr Cope, KSdd. While the taxonomic status of these populatitms has been c]uestioned for 
many decades, a tliori>iii;h ie\ it'w ot the B. coccifer complex never has been presented. Based on evidence from 
external morphology and a partial molecular data-set, we conclude that this complex consists minimally of six 
species. Herein, we recognize B. ivicifcr Cope, 1866, B. cychidcii Lynch and Smith, 1966, and B. ibanai Stuart, 
1954, and describe three new species. 

Key Words: Bufonidae; Bufo coccifcr, Biifo cycladcn, Biifo ilnirrai, Biifo /'/s/mhhs, Bufo porlcri, Bitfo signifer; 
Mesoamerica; taxonom\'. 



KESUMEN Muchas poblaciones de sapos que se encuentian cntre el oeste-central de Mexico y Panama 
han sido referidas como Bufo coccifcr Cope 1866. Aunc]ue el estado taxoncimico de estas poblaciones ha sido 
cuestionado por muchas decadas, nunca se ha presentado una revision completa del complejo B. coccifcr. En 
base a c\ idcncia do mortologi'a external y una base de datos moleculares parcial, concluimos que este complejo 
consiste de al menos seis especies. En este trabajo, reconocemos B. coccifcr Cope, 1866, B. cycladcn Lynch and 
Smith, 19h6, y B. ilnirrai Stuart, 1934, y describimos tres especies nuevas. 

Palabraf Clave: Bufonidae; Bufo coccifcr, Bufo cycladcn, Bufo ibnrrai, Bufo pisnmus, Bufo portcri, Bufo signifer; 
Mesoamerica; taxonomfa. 

INTRODUCTION 



The taxon Bufo coccifcr Cope, 1866 currently is applied 
to toads that are distributed allopatrically in five regions 
of Mesoamerica (Pig. I): (1) the Tepalcatepec Valley, 
Michoacan, Mexico; (2) the Pacific slope of the Sierra 
Madre del Sur in Guerrero, Mexico; (3) the southern side 
of the Isthmus of Tehuantepec, Oaxaca, Mexico; (4) nearly 
the entire Pacific versant from northwestern Guatemala 
to northwestern Costa Rica; and (5) western Panama. 
Additional records exist from the Atlantic versants of 
Honduras and Nicaragua. The distribution of these 
populations approximately matches the distribution 
of low elevation tropical dry forest (Rzedowski, 
1994; Campbell, 1999) in these regions. However, the 



Rica: Cope, 1866:130) and suggested that the holotype 
likely originated from somewhere on the Meseta Central 
of Costa Rica; this referral was supported by Savage (1974). 
Porter (1963) provided a range map and general diagnosis 
of B. coccifcr Porter (1965) discussed the distribution of 
the species in more detail, observing that the populations 
in Michoacan, Mexico, and Oaxaca, Mexico, evidently 
are allopatric — with a hiatus of approximately 140 km 
between records from Oaxaca, Mexico, and records from 
southeastern Guatemala. Stuart (1954a:20) referred to "a 
chain cif Bufo coccifcr-Wke toads" distributed through the 
subhumid habitats of Central America, and specifically 
noted undescribed species in central Guatemala, and 



population on the Pacific slope of the Sierra Madre del Sur another in Guerrero and the Isthmus of Tehuantepec 



is an exception to this generality; these toads are found at 
moderate elevations (ca. 1000 m) in relatively wet pine- 
oak forest. (See Campbell and Duellman, 2000, for habitat 
description.) Other records from more mesic, upland 
habitats include the mountains of central Honduras 
(McCranie and Wilson, 2002). An additional specie.s, B. 
ibarrai Stuart, frequently has been assigned to the Bufo 
coccifcr Group (e.g.. Frost, 1985). The taxonomic status 
and distribution of B. ibarrai was reviewed by Mendelson 
(2001); this species occurs primarily in upland pine-oak 
habitats in Guatemala and is reported here lor the first 
time in an adjacent region of Honduras. 

With reference to the broad distribution ot lUdo coccifcr, 
it has been suggested many times that these wirious 
populations likely are not conspi-citic (Stuart, 1954a, 
1963; Duellman, 1960; Porter, 1963, 1965; Zweifel, 1965; 
McDiarmid .\nL\ I oster, 1981; Mendelson, 2001; McCranie 
and Wilson, 2002). Dunn and Stuart (1951) commented on 



region of Mexico. Stuart (1954b) subsequently described 
the populations in the upland pine-oak zone of Guatemala 
as B. ibarrai. Lynch and Fugler (1965) reported B. ibarrai 
from Honduras. Subsequently, the taxon has had a long 
period of uncertain status, typically being confused with 
B. coccifcr (e.g., comments by J. A. Campbell in Frost, 
1985:41). Meyer and Wilson (1971) placed B. ibarrai in 
the synonymy of B. coccifcr. However, Mendelson (2001) 
recognized B. ibarrai as a ciistinct species, described the 
tadpole, and prcnided a new diagnosis and range map 
of records tiom Guatemala; also, he suggested that 
the species may be endemic to Guatemala. Based on 
examination of specimens from Honduras, McCranie 
and Wilson (2002) suggested that B. ibarrai does not 
occur in that country. Lynch and Smith (1966) referred all 
populations in Mexico to the new taxon Bufo cycladcn and 
cited differences in the ad\'ertisemenl call (tidi' Poitei; 
1965) and ecology of the Mexican and Central American 



thestatusof the type locality for B. coccifcr (" hrnha" Costa populations. I lo\ve\'er, the morphologv of this t 



ixon was 



Toads oi- iiii- Bcro coccifir Comi'LLX 




K ilometers 



100 



Fij;, 1 , A m>ip of MfsoaniL'iiLM, sluuving thu genoriili/ed di^li ibutioiis of fo.ids ivtc-ned to Biifo locci/ci soiimi Kito, .ind of /x ibaiiai (as recognizud 
hi'iein). 



diagnosed poorly, with respect to B. ibnrrai (see Mendelson, 
2001 ) cind other popiilatiotis referable to B. coccifcr (Porter, 
1967). Porter (1%7) criticized Lynch and Smith (1966) for 
not supporting their claitns of timrphological differences 
atnong the populations atid also claimed that there are 
no real ecological differences among the populations 
considered; Porter proposed this ta\on to be a mniicii 
diibiiuii. The taxonotnic cofifiision surroiuidifig B. c\/cliu1cii 
is evident iti the fact that the taxon appears Ofi some recent 
checklists (e.g., Frost, 1985), but not oti others (e.g., Flores- 
Villela, 1993; Catnpbell, 1999). 

Se\eral authors have described the male ad \ertisefnent 
calls from various areas. Porter (1964, 1965) described the 
ad\'ertisement calls of B. cociifcr from the Tehuantepec 
region of Oaxaca, Mexico, in addition to those of toacHs 
he referred to B. coaifcr recorded in Guerrero, Mexico, 
and several populations in Central Atnerica (El Salvador, 
Honduras, Nicaragua, and Costa Rica; Porter, 1965). The 



various descriptions and comparisons of advertisement 
calls presented by Porter (e.g., 1964, 1965) are difficult 
to interpret because he provided no voucher nufiibers 
for recorded specimens, and the detail of his locality 
information is inconsistent. When tracking down specific 
localitv data provided by K. R. Porter, we found it useful 
to refer to recording station descriptions presented in 
his original, unpublished dissertation (Porter, 1962:table 
1). Zweifel (1965) first reported the presence of toads he 
referred to B. coccifcr in Panama and demonstrated that the 
advertisement call of this population differed from those 
of both the Mexican (= Isthtnus of Tehuantepec region of 
Oaxaca) and other Central Americati recordings that were 
published by Porter (1964). McDiarmid and Foster (1981) 
described additional advertisement calls recorded in Costa 
Rica, and compared them to those reported from Mexico 
and Central America (Porter, 1965) and from Panama 
(Zwiefel, 1965); there are substantial differences in pulse 



SciENTiiic Papi:rs. Natlrai. Hisiorv MusiiUM.Tiiii UNi\i;Rsir-t oi Kansas 



labk' 1. Ciimporison ot i\ingi"s or moiins ot c\ills of the Hufo coccifcr ciimplex Ironi dlllcrcnt populations. Tlic identity ot the toads recorded 
from Honduras (Porter, 1965) are unknown because the original paper provides neither reference to voucher specimens nor a detailed localilv- 
description; thiTc ,ire at least three species in this group in ?londuras. 





.\o. individuals 


Duration 


Frequency 


Pulse Rate 




Species 


Location 


recorded 


(second) 


(Hertz) 


(per second) 


Source 


identity 


Mexico: Guerrero 


1(6 calls) 


mean =5.7 


mean =2800 


mean =120 


Porter, 1965 


B. cycladcn 


Mexico: Oaxaca 


18 


1.4-6.4 


2800-3350 


97-115 


Porter, 1964, 1965 


B. coccifcr 


El Salvador 


14 


3.0-16.3 


2200-2700 


80-101 


Porter, 1965 


B. coccifcr 


Guatemala 


7 


4.2-5.8 


1650-1800 


48-61 


Porter, 1966 


B. ilmrrai 


Honduras 


2 


5.3-8.8 


2200-2500 


80-96 


Porter, 1965 


unknown 


Nicaragua 


4 


6.2-10.9 


2300-2400 


91-100 


Porter, 1965 


B. coccifcr 


Costa Rica 


4 


1.5-10.0 


2300 


90-95 


Porter, 1965 


B, coccifcr 


C\>sta Rica 


1 


1.2-5.7 


2231-2539 


90-105 


McDiarmid & 
Foster, 1981 


B. coccifcr 


Panama 


1 


8.0-17.0 


2100-2600 


66 


Zweifel, 1965 


B. sigtiifer 



rates and dominant frec|Liencies among these samples. 
Porter (1966) described the ad\'ertisement call of B. ibiinui. 
The data reported bv I^orter, Zweifel, and McDiarmid and 
Foster are presented in Table 1 and summarized in Figure 
2. We are unaware of any recordings of populations of 6. 
cf. coccifcr from Guatemala or Michoacan, Mexico. 

The calls of male toads from the Sierra Madre del Sur 
of Guerrero and the Tehuantepec region of Oaxaca have 



higher dominant frequencies than calls of all other samples. 
The call of the toad in Guerrero also has a higher pulse rate 
than all other reported calls. Note, however, that the data 
from Guerrero are based on a recording of six separate 
calls by the same individual. These data were used, in 
part, by Lynch and Smith (1966) to justify recognition of all 
Mexican populations (including populations in Michoacan, 
Guerrero, and Oaxaca) as a species (B. cyclndcii) distinct 



Dominant Frequency 



Guerrero 
Oaxaca 

Central America 
Panama 
B. ibarrai 



1600 



2500 
Hz 



~1 
3400 



Pulse Rate 



Guerrero 
Oaxaca 

Central America 
Panama 
B. ibarrai 



40 



I 

80 
pulse/sec 



120 



Fig. 2. Cirapliical representation ol Irequencies ami pulse rate's ot advertisement calls Ironi toads ul Ihe Biito coccifcr complex Ironi dilterent 
regions of Mesoamerica; information shown here are a graphical representation of data presented m l.ible I IXila from "Cenlral America" here 
includes range of values representing samples from El Salvador, Nicaragua, and Costa Rica. 



Toads of thi-; Bufo cocciii-r Comi'i k.\ 



expressed or implied consensus of these authors is 
that a thorough revision of the Biifo coccifcr complex is 
long overdue, in tliis paper we use data from external 
morphology and male advertisement calls (previously 
published) to delimit species boundaries amting samples 
referable to the widespread taxon B. coccifcr. We also present 
a preliminary assessment of phvU>genetic relationships 
among the species in the group, based on mtDNA data 
from several of the species. 

Acknowledgments 

We are grateful to the following inclividuals for 
providing assistance in the field, laboratory, and library: W. 
Duellman, S. Cotte, K. Lips, M. Acevedo, E. Greenbaum, 
J. R. McCranie, J. Malone, D. Laurencio, M. Sasa-Marin, 
M. Ryan, M. Forstner, T. Reeder, J. Campbell, E. Smith, 
C. Franklin, and R. CJutberlet. Adele Cutler provided 
statistical acivice and assistance. Michelle Koo graciously 
provided access and assistance with mapping software at 
the California Academy of Sciences. We benefited greatly 
from memories, observations, and notes drawn from the 
considerable fieki experiences of J. Campbell, C. Myers, 
j. Sa\'age, W. Duellman, anci R. Zweifel. Permission and 
help with photographs was provided by M. Sasa-Marin, 
R. Zweifel, and J. Simmons. Loans of critical specimens, 
for extended periods of time, are gratefully acknowledged 
from the curators and collection managers of the 
following institvitions: AMNH, CAS, FMNH, TCWC, 
UTA, TNHC, ANSP, UIMNH, USNM, KU, UMMZ, 
MVZ, LACM, and LSUMZ. Some of the fieldwork for 
this project was supported by funds from The National 
Geographic Society and this research was conducted in 
direct association with The Research Analysis Network 
for Neotropical Amphibians (RANA; NSF-DEB 0130273). 
Comments on the manuscript were provicied by the 
students and faculty of the USU Herpetology Group, L. 
frustration in delimiting species in this complex, which Trueb, J. Pramuk, and W, Duellman. Additional help and 
is characterized bv advertisement calls that vary among favors were provided by J. Meik and G. Schneider, 
populations and subtle morphological differences. An 

MATERIAL AND METHODS 



from B. coccifcr in Central America. A call described by 
Zweifel (1465) from Panama differs dramatically from all 
others b\' having a much lower pulse rate. The relationship 
between the calls from Oaxaca and Central America is 
interesting in that they have distinctly different ranges 
of dominant frequencies and somewhat different (but 
o\'erlapping) ranges of pulse rates. However, we note 
that the Central America data do not include recordings 
from Guatemala — the closest geographic region southeast 
of the Oaxacan population. Recordings from Guatemala 
may tiiminish the apparent distinctit)n between the 
advertisement calls from the Tehuantepec region of Oaxaca 
and Central America. Porter (1966) described the call of B. 
ibarrni from Guatemala; this species differs from all other 
populations in the B. coccifcr complex by having a very low 
frequency and pulse rate (Table 1). 

In their description of Bufo ci/chhicii, Lynch and Smith 
(1966) placed great diagnostic importance on variation in 
calls between Mexican and Central American populations. 
We note that their summary of calls from Mexico was a 
composite of data from both Guerrero anti the Tehuantepec 
region of Oaxaca (data published by Porter, 1964, 1965; Table 
1 ). VVe have concluded that these toads from Guerrero are 
not conspecific with those from Oaxaca (discussed below). 
Gergus et al. (1997) described call variation among toads of 
the B. Diicroscnpliufi complex; they reported wide variation 
in some parameters and an overall pattern of apparent 
plesiomorphic similarity among allopatric species. There is 
no reason to expect drastic differences in mate-recognition 
systems among allopatric members of a complex of closely 
related anurans with similar natural histories (Gergus et 
al., 1997). Nevertheless, there are apparent differences in 
the advertisement calls of toads of the B. coccifcr complex 
that suggest diagnostic differences in the mate-recognition 
systems of these allopatric populations. 

All previous workers (above citations) have expressed 



General terminology and measurements are those of 
Mendelson (1997). Adult males were identified by the 
presence of vocal slits and nuptial excrescences; large 
individuals lacking these characters were presumed to be 
adult females. If sex could not be determined externallv, 
it was verified by direct observation of the gonads. 
Foot-webbing formulae follow the system of Savage 
and Heyer (1967), as modified by Myers and Duellman 
(1982) and Savage and Heyer (1997). The general format 
of the descriptions and diagnoses is slightly modified 
from that of Mendelson (2001). Museum codes are those 
proposed by Leviton et al. (1985). We follow Tyler et al. 



(2001) in OLu- usage of the term "parotoid gland." Data 
from advertisement calls were taken from the literatiuv 
(referenced below). 

MORPHOMI^TRIC DaIA 

The following measurements were taken from adult 
specimens: snout-vent length (SVL); head length (HL); 
head width (HW); tibia length (TIB); foot length (FL); width 
of tympanum (TYM); length of parotoid gland (PARE); 
maximum width of parotoid gland (PARW); and length of 
supratympanic crest (SPTYM). These variables represent 
repeatable morphological landmarks and were measured 



6 



Scientific Papers, Natural History Museum. The University oi- Kansas 



with digital calipers and iDiinded tt) the nearest 0.1 mm. 
Because of the paucity of large series of females from many 
localities, all morphometric analyses are based only on 
adult males. Many of the data for Biifo ibnrnii are the same 
presented by Mendelson (2001), but supplemented with 
additional material frc)m both Guatemala and Honduras. 
We conducteci a Principal Components Analysis 
(PCA) using the covariance matrix on log-transformed 
morphometric measurements from 324 adult male toads; 
this data-set included specimens representing all species 
and geographic regions of the distribution of the Bufo coccifcr 
complex. We also performed a PCA on the residuals of the 
latter seven variables derived from a regression analysis 
(using SVL as the independent variable). This type of PCA 
removes the effect of size, and displays general variation 
in shape and proportion among the specimens (Good and 
Wake, 1992). We performed a Linear Discriminant Function 
Analysis (LDA) on the same data-set, and used residuals 
of variables regressed on SVL and log-transformed SVL. 
In this analysis, a priori groupings corresponded to the 
species recognized in this paper (Accounts of Species, 
below). Statistical analyses were performecl using SAS 
and Minitab software. 

Molecular Data 

We conducted molecular analyses on ail available 
samples, which inckided five specimens of Bufo coccifer (4 
from Central America, 1 from Guerrero, Mexico) and two 
specimens of B. ibnrrai. Additionally, one specimen of each 
of the following species was included as outgroup taxa: 
B. conifcrus, B. valliceps, and B. mnrijius. The selection of 
these outgroup taxa was based on preliminary analyses of 
a data set containing approximately 25 species of Central 
American bufonids (Mendelson and Mulcahy, in prep.). 
Sections of tiie mitochondrial genes cytochrome-b (cyt-b) 
and 16S were used in the molecular analyses. Isolation and 
PCK amplification of the mitochondrial DNA (mtDNA) 
genes were performed exactly as described in Mulcahy 



and Mendelson (2000), which includes primer information 
and amplification prt)files. Products from the PCR were 
amplified and sec]uenced in both directions using BigDye™ 
Terminator Cycle Set]uencing Ready Reaction Kit (Applied 
Biosystems Part No. 4303152); we used the same PCR 
primers and standard sec]uence-reaction profile on a Perkin 
Elmer GeneAmp 2400 cycle sequencer. Cleaned sequences 
were then run on an ABI 377 automated sequencer by 
DGM at the Biology Department at Utah State University. 
Sequence comparisons and alignment were conducted 
with Sequencher 3.1. 

Phylogenetic analyses of the mtDNA sequences 
were conducted using PAUP* 4.0b8a (Swofford, 2002). 
A partition-homogeneity test of 100 replicates was 
implemented in PAUP (using default parsimony settings, 
with the exception of addition-sequence being random 
with 100 replicates) between cyt-b and 16S gene regions to 
determine whether or not the two genes yielded conflicting 
results. Maximum-parsimony analyses (MP) were 
performed on each gene separately and both combined. 
The program Modeltest (Posada and Crandall, 1998) was 
used to evaluate the best Maximum-likelihood model, using 
the Hierarchical Likeliht)od Ratio Tests (hLRTs) criterion. 
A Maximum-likelihood (ML) analysis was then conducted 
using the model and settings based on the hLRTs results. 
Because of the limited number of taxa in the phylogenetic 
analyses, an exhaustive search was possible, and used in 
the MP analysis, while the ML analysis required a heuristic 
search algorithm. Gaps were treated as "missing data," 
with characters-state optimization set at ACCTRAN. 
Branch support was assessed bv nonparametric bootstrap 
analyses using 1000 replicates of full heuristic searches, 
with 100 random additions at each replicate, under MP and 
100 replicates of full heuristic searches, with 10 random 
additions under the ML criteria. Decay indices (Bremer, 
1994) were also measured under the parsimony analysis 
using the program TreeRot (Sorenson, 1996). 



ACCOUNTS OF SPECIES 



We used data from adyertisement calls, nu)rphometry, 
external morphology, and DNA sequences to examine 
variation among samples of the Bufo coccifcr complex. 
These data are consistent with the "chain of Bufo 
coccifer-Vikc toads" distributed through the subhinnid 
habitats of Central America that Stuart ( 1954a) envisioned. 
Evaluation of our data with respect to the Evolutionary 
Species Concept (sensu Wiley, 1978; Frost and Hillis, 
1990) supports recognition of six species in this complex. 
Our proposed taxonomy reflects the sentiments of the 
many authors (e.g., Stuart, 1954a, 1963; Duellman, 1960; 
Porter, 1963, 1965; Zweifel, 1965; McDiarmid and Foster, 
1981; Mendelson, 2001; McCranie and Wilson, 2002) who 



have dealt with these toads during the last five decades. 
We provide species accounts and diagnoses for B. coccifer, 
B. ci/cladcn, and B. ibnrnii, and describe three new species 
from this complex; photographs of these species in life are 
presented in Figure 3. 

Bufo coccifcr Cope 
Figs. 3-5 

Bufo coccifcr Cope, 1866:00. Hiil.it\pc: USNM 6490. Type locality: 
"Arriba" Costa Rica. 

lUifo coccifcr — Dunn and Emlen, 1932; Kellogg, 19,12; Hartweg and 
(.liner, 1940; Dunn and Stuart, 1951; Smith and Taylor, 1948 [in part 
Mertens, 19.S2; Taylor, 1932; Stuart, 1954b; Rand, 1957; Duellman, 1960 
I'orter, 1963 |in part); Stuart, 1963; Porter, 1964; Porter, 1965 (in part] 



Toads of thh Bufo coccifer Complex 




Fig. 3. Species of the Bi{fo coccifer complex in life, clockwise from upper left: B. coccifer from Santa Rosa, Costa Rica (adult male: photograph 
hy Andrea Bernecker); B. cyclndcii from Guerrero, Mexico (adult male, UTA-JRM 4607; photograph by ]. R. Mendelson); B. ibarrni from Baja Verapaz, 
Guatemala (female, KU 186304; photograph by J. A. Campbell); B. pifimiiif from Michoacan, Mexico (adult male, from UMMZ series; photograph 
bv W. E. Duellman); B. portcri from Francisco Morazan, Honduras (subadult female, KU 103220; photograph bv W. E. Duellman); B. si\;uifcr (adult 
female, AMNH 69625; photograph by R. G. Zweifel). 



Sfii:Nrii i( P,\i'i-.ks. Nvn RAi, lIisroK^ Mii.si-;um,Thk Univhrsii'i oi- Kansas 



Lvnch and Fuglor, 1965; Porler and Porter, 1967; Meyer and Wilson, 1971 
|in part); Villa, 1972; Savage, 1974; McDiarmid and Foster, 1981; Villa, 
1983; Frost, 1985 |in part]; Savage and Villa, 1986; Villa et al., 1988; |in 
parti; Campbell and Vannini, 1989 [in part]; Flores-Villela, 1993 |in part]; 
Campbell, 1999 |in part]; Kiihler, 1999; Mendelson, 2001; Frost, 21)03 |in 
parti; McCranie and Wilson, 2002 |in part]; Savage, 2002. 

Biifo cifcliuleii — Lynch and Smith, 1966 [in part, for reference to 
specimens from Tehuantepec region of Oaxaca, Mexico]; Frost, 1985 ]in 
part]; Frost, 2003 ]in part], 

Biifo iharrai — Lynch and I'ligler, 1965 [in part]. 

Bufo vatlicepf micwli^ — Werner, 1896 (synonyiiu' b\' Mendelson, 
2001). 

Diagnosis. — A medium to large species of Bnfo 
{males to 62 mm SVL; females to 82 mm SVL) having 
the following combination of characters: (1) tympanum 
e\'iclent externally, about 35-45' V' diameter of orbit in males 
and females; (2) cantlial, supraorbital, supratvmpanic, 
postorbital, preorbital, pret\'mpanic, parietal, and 



siipralabial crests present; (3) cranial crests well dexeloped, 
i(.)biist, except parietal mav be thin, low, or absent in some 
specimens; (4) tibia short, about 35% SVL; (5) feet short, 
about 35% SVL; (6) dorsal tubercles small to medium 
sized, elevated, rounded, scattered relatively sparsely 
on middorsal, dorsolateral, and lateral regions of bod\' 
beci)ming more denseh' arranged and distinctly spinose 
laterally in specimens from most regions; (7) yentral 
tubercles granular, smooth, or with spinose apices; (8) 
lateral descending row of enlarged tubercles absent; (9) skin 
texture not sexually dinn)rphic; (U)) \ocal slit unilateral in 
male; (11) in. iiitcrln/oidciii^ poorly diflerentiated from ;;;. 
iiitciiiiaiidibiilnris, but differentiated posteriorh' forming a 
large, unilobed vocal sac with heavy black pigmentation; 
(12) snout rounded in lateral profile, pointed in dorsal 
aspect; (13) parotoid glands round to oxoid and large, about 




Fig. 4. Dorsal and ventral aspiTts ol representalise adult spi-ciniens of Hufo coctifcr from Ketalhiileu, C.iiatcniala (male, letl: UT.'\ .A-2,5821, SVL 
= 57.0 mm; female, right: UTA A-29025, SVL = 75.0 mm) and B. ci/chulcn Irom t.iierrero, Mexico (male, lelt: L'MM/ 1 1 5357 ] WHO 92751, SVL = 53.8 
mm; female, right: UMMZ 119270 ]WKD 1.3425], SVL = 61.4 mm). 



ToAUs oi- iiii-; Brio coccii i:k Comi'lilX 





Fig. 5. L.itei mI .ispccts ot tlif hiMd'i i.)l adult nulf >pf cimens of Biifo 
coccifer (upper: KU hS4(l(l) .ind B. cycln'icu (lower: KU 97434). Scale bars 
= 1 cm. 

1.0-1.5 times size of eyelid; (14) skin between cranial crests 
on top of head with few to many scattered, low, rounded 
tubercles; (15) ventral coloration whitish cream, sometimes 
with some degree of diffuse dark pigmentation, stunetimes 
in the form of diffuse punctuations. 

Specimens referable to Biifo coccifer vary considerably 
in size, shape, coloration, skin texture, and parameters of 
the advertisement call across the range of the species. Biifo 
coccifer differs from B. ci/cladeii by: being larger (males of B. 
cyclndcii to 54 mm SVL, females to 62 mm); having dorsal 
tubercles that are relatively small and scattered about the 
dorsal surfaces, rounded on dorsum, becoming spinose 
laterally in most specimens (tubercles large, elevated, 
and densely arranged in B. cycliiden, rounded dorsally, 
becoming very spinose laterally); ventral tubercles that 
are smoothly granular or with small, spinose apices 
(ventral tubercles in /-!. c\/clih1eii with distinct, large 
spinose apices); relati\el\' well de\eloped parietal crests 



that rarely are absent (parietal crest weakly de\'eloped in 
B. cyclndeu, sometimes absent); and an advertisement call 
with a lower pulse rate (Table 1; Fig. 2). Biifo coccifer can be 
distinguished from B. pisiumts by: being larger (males of 
B. yisiuiiiifi to 51 mm SVL, females to 62 mm); and having 
skin tubercles that are overall larger (all tubercles minute 
in B. pi:^iuiiiis). Biifo coccifer may be distinguished from B. 
signifer by having: no, iir few, relatively indistinct dark 
brown markings on the venter (that of B. i^igiiifcr boldly 
marked with a marbled pattern); relatively thicker and 
higher cranial crests (all crests relatively low and thin 
in B. signifer); and an advertisement call with a higher 
frec^uency and pulse rate (Table 1; Fig. 2). Biifo coccifer 
differs from /->. ihnrnii by: being smaller (males of B. 
ibarrai to S2 mm SVL, females to 94 mm); having rounded 
to subovoid parotoid glands about 1.0-1.5 times size of 
eyelid (parotoid glands distinctly ovoid, about 1.5-2.0 
times size of eyelid in B. ibarrni); and males usually with 
dorsal tubercles becoming spincise laterally and females 
with rounded tubercles middorsally (dorsal tubercles of 
males of B. ibarrai rounded laterally and females with 
spinose middorsal tubercles). Bufo coccifer differs from 
B. porteri by having: sharply spinose lateral tubercles in 
males (rounded in males of 6. porteri); and a relatively 
thin supratympanic crest (large, bulbous in B. porteri). 
Morphometric variation is summarized in Table 2. 

Distribution and Ecology. — Bufo coccifer occurs along 
the Pacific versant of Mesoamerica, from the Guatemala- 
Mexico border to the Guanacaste region of Costa Rica 
(Fig. 6). Additional records exist from interior valleys on 
the Atlantic versant of Honduras and from the Atlantic 
coastal regions of Honduras and Nicaragua. McCranie and 
Wilson (2002) erroneously reported this species from the 
Atlantic versant of Guatemala; there are no records from 
this region. An apparently isolated population occurs in 
the southwestern region of the Isthmus of Tehuantepec, 
Oaxaca, Mexico. This species ranges from sea level to 1435 m 
(Savage, 2002; record from Cartago, Costa Rica) and occurs 
in a variety of tropical dry forests and savanna habitats. 
Duellman (I960) provided photographs of the habitat 
and some natural history observations from the Isthmus 
of Tehuantepec. McDiarmid and Foster (19cSI) described 
the reproductive biology of a population in northwestern 
Costa Rica. The ecological notes provided by McCranie 
and Wilson (2002) for Hondiu'an populations represent 
mixed observations pertaining to B. coccifer, B. ibarrni, and 
B. porteri. Savage (2002) provided a concise summary of 
the ecology of this species, based on observations from 
Costa Rica. 

Tadpole. — McDiarmid and Foster (1981) and Savage 
(2002) described tadpoles from Costa Rica and provided 
illustrations. 



10 



Scientific Papi-ks. Nau kai History MLisi-.uM.THi-. Univi-ksha oi Kansas 



Table 2. Morphometric variation among males and femak's from of the Biifo ccccifcr complex; mean + SD is presented above the range (in mm). 
Data for B. ibarrai taken, in part, from Mendelson (21)01 ). 





S. cocciff) 


B. cydadcn 


B. ibarrai 


/' yi'-nniu-^ 


/ ■' 


:: <.-:lfcr 


Males 


u = 130 


n ^ 311 


u ^ 79 


n = 2l 


It =43 


» = 21 


SVL 


51.4 ±4.8 


51.1 ±2.2 


62.2 ±9.1 


45.9 ± 3.2 


49.6 ± 4.6 


51.2 ±64 




41.0-64.4 


47.4-54.2 


42.5-82.4 


38.5-51.4 


40.2-58.7 


42.0-64.1 


Tibia length 


18.3 ±1.9 


18.4 ±0.9 


24.1 ± 3.6 


15.9 ±1.4 


18,4 ±1.8 


17.9 ± 2.3 




13.3-22.5 


16.6-20.1 


15.7-31.8 


13.0-18.0 


15.2-21.7 


15.0-23.4 


Foot length 


18.1 ± 1.9 


18.3 ±0.7 


24.6 ± 3.8 


16.9 ±1.3 


18.8 ±1.8 


18.2 ± 2.3 




13.1-23.9 


16.9-19.8 


16.3-33.4 


14.8-19.2 


15.7-22.7 


13.7-23.1 


Head length 


17.8 ± 1.8 


18.0 ±0.9 


20.8 ± 2.8 


15.1 ± 1.0 


16,7 ±1.4 


174 ±2.0 




13.6-21.6 


16.7-20.9 


15.3-26.5 


12.9-17.1 


14.0-20.0 


14.7-22.2 


Head width 


20.5 ± 2.4 


20.1 ±0.9 


23.9 + 3.1 


17.3+1.4 


18.9 ±1.6 


19.8 ± 24 




15.9-27.8 


17.6-21.8 


16.2-30.8 


15.1-20.2 


15.7-23.0 


16.6-24.9 


Tympanum 


2.3 ± 0.5 


2.7 ± 0.3 


3.7 ± 0.7 


2.3 ± 0.3 


3.0 ± 0.4 


3.1 ± 0.4 




1.3-3.7 


2.1-3.3 


2.7-5.3 


1.8-2.6 


2.2-3.5 


2.5-3.8 


Supratympanic 


3.2 + 0.5 


2.7 ± 0.3 


3.5 ± 0.5 


2.6 ± 0.3 


3.1 ±0.3 


3.0 ± 04 


crest 


2.0-4.4 


2.2-3.3 


2.3-5.2 


2.2-3.2 


2.3-3.7 


2.1-3.8 


Parotoid length 


6.3 ±1.0 


6.9 + 0.5 


7.8+1.5 


5.4 ± 0.6 


6.3 ± 0.9 


6.3 ±1.2 




4.3-8.7 


6.0-4.8 


3.2-11.0 


4.1-6.5 


5.0-9.0 


2.8-8.8 


Pan>toid width 


6.6 ±1.3 


6.1 ±0.6 


5.7 ± 0.9 


5.3 ±0.5 


5.1 ±0.7 


6.0 ± 0.8 




3.7-9.5 


5.0-7.2 


3.5-7.4 


4.5-6.2 


3.8-6.7 


4.6-7.7 


Females 


It = 25 


n =6 


n = 61 


» = 5 


// = 14 


11 = 11 


SVL 


67.7 ± 7.6 


59.8 + 2.8 


78.4 ± 7.6 


59.1 ± 3.3 


64.5 ± 5.8 


67.2 ± 8.3 




58.1-82.4 


54.6-62.9 


60.8-94.4 


55.4-62.2 


56.4-73.0 


54.5-77.0 


Tibia length 


22.2 ± 2.4 


20.0 ±1.6 


28.7 ± 3.0 


18.7 ±1.1 


77 5 + 2.2 


23.0 ± 3.4 




18.8-28.1 


18.3-22.3 


20.2-32.8 


18.0-20.6 


18.7-27.2 


17.7-26.5 


Fool length 


21.9 ±2.2 


20.4 ±1,0 


28.7 ± 3.7 


20.3 ± 1.3 


22.5 ±1.9 


23.1 ±3.7 




17.9-27.2 


19.4-22.1 


19.6-34.1 


18.8-21.7 


19.6-26.5 


17.0-26.8 


Head length 


?? ? ± 2.2 


20.7 ±7.6 


25.2 ±2.1 


18.8 ±0.9 


20.9 ± 2.0 


22.6 ± 2.5 




19.2-26.8 


18.9-22.8 


19.8-28.5 


17.6-19.7 


18.0-25.9 


18.4-25.9 


Head length 


25.5 ±2.5 


23.0 ±1.5 


29.5 ± 2.8 


21.9 ±1.2 


23.9 ±1.9 


26.1 ±34 




22.5-31.5 


21.1-24.4 


21.5-33.3 


20.6-23.7 


21.5-28.3 


20.9-29.4 


Tympanum 


3.1 ±0.6 


2.8 ± 0.5 


4.1 ±0.5 


3.2 ±0.1 


3.5 ± 0.5 


3.6 ± 0.4 




2.1-4.2 


2.2-3.5 


3.0-5.4 


3.1-3.4 


2.6-4.5 


3.1-4.3 


Supratvnipanic 


4.3 ± 0.8 


3.1+0.2 


4.5 ± 0.7 


3.4 + 0.3 


3.9 ± 0.6 


4.0 ± 0.7 


crest 


2.9-6.2 


2.8-3.4 


2.9-6.0 


3.1-3.8 


3.0-5.2 


3.1-5.2 


Parotoid length 


7.4 + 1.3 


7.6 + 0.8 


8.4 ±1.5 


6.8 ±1.0 


8.3 ± 08 


7.8 ±1.3 




5.4-9.9 


6.5-8.5 


3.0-11.7 


5.9-8.4 


6.8-9.9 


5.5-9.9 


Parotoid width 


7.4 + 1.1 


6.9 ±0.8 


6.3 ± 0.9 


6.6+ 1.0 


6.6 + 0.4 


7.7 ±1.0 




■i 7 1114 


6.2-8.4 


4.1-8.2 


■^ (. ~ s 


b 1 - 2 


5.4-9.0 



Remarks. — Ci)k>r photographs ot this species appear 
in Leenders (2()01:pl. 7), Savage (20n2:pl. 77), and Villa 
(1972:fig. 57). Blair's (1963) reference to Bufo coccifcr in his 
discussion of the evolution of North American bufonieis 
is unclear because he does not give the provenance of 
the toads he analyzed. Lynch and Smith (1966) referred 
specimens from the Tehuantepec region of Oaxaca, Mexico 
to B. ci/clndcii; however, this designation was not followed 
by nn)st subsequent authors (Porter, 1967). Toads from the 
Tehuantepec region are distinct and aiiopatric, with respect 
to the toads we refer to B. cyclndcii on tlu' slopes ol tiie Sierra 
Madre del Sur in Guerrero and western Oaxaca. 

There are no records of Bufo coccifcr from the Pacific 
Coastal Plain of Chiapas, Mexico. Despite tiie lack oi 
records in this region, we note that much of the region 



seems to be suitable subiiumid habitat. Subhumid forest 
and scrub habitats extend along the coastal plain here, but 
grade into more humici forests along the base of Volcan 
Tacanci (Johnson, 1989; Rzedowski, 1994). Although the 
lone specimen from Tapachula appears to have been 
found in a humid region, we note that the historical humid 
forests (W. E. [3uellman, pers. comm.) along the coastal 
margin of Volcan Tacana have been completely converted 
to crop and pasture lands (pers. obs.). Further fieldwork 
along the Pacific Coastal Plain of Chiapas is warranted 
to \eiif\ the allopatr\' of the population of B. coccifcr in 
the Islhnnis ot Ti'huanatepec, with respect to conspecifics 
in Ciuatemala. Johnson (19S9) implied that B. coccifcr 
occurs along the Pacific Coastal Plain of Chiapas, perhaps 
based on records from Soconiisco, Chiapas, that were 



Toads oi nil-; Brio C(kxii ii< C()m1'L[:.\ 



11 




[■ij;. li- \Lip of Central AmcriLM showing the geogr.iphie distribution of Bufo coccifcr. B. ibarnii .ind B. poi Ifi i. 



listed by Porter (1963); these records actually represent 
misidentified specimens referable to 6. imlliccpf. The single 
specimen from Tapachula, Chiapas (MVZ 177594), is a 
recently metamorphosed specirnen tiiat originally was 
misidentified as B. luetkeiii. This record seems to be the 
source of information that resulted in B. luetkenibis'mg listed 
incorrectly as part of the Mexican herpetofauna (Johnson, 
1989; Flores-Villela, 1993)— B. luctkcni does not occur in 
Mexico. Johnson (1990) correctly stated that records of B. 
coccifcr from the Grijalva Valley of Chiapas, Mexico, were 
based on misidentified specimens. 

Biifo ci/clndcii Lynch and Smith 
Figs. 3-5 

Bufo cydaden Lynch and Smith, 1966:19. Holotype: UIMNH 57142. 
Type loCiiHty; 3 mi |4.8 km] S Putio de Guerrero, Oaxaea, Mexico. 



Bufo Li/cliidcii — Porter, 19h7 (decKired luvucii (luluuiii\; Frost, 1985 |in 
part]; Frost, 2003 |in part]. 

Bufo aicn/I-i— Smith and Ta\ lor, I94S |in part]; Porter, 196,3, 1965 |in 
part]; Villa et al.. 19SS |in part): Flores-Villela, 1993 |in part); Campbell, 
1999 [in part]. 

Diagnosis. — A small species of Bufo (males to 54 
mm SVL; females to 62 mm SVL), having the following 
combinationofcharacters:(l)tympanumevidentexternally, 
about 40-50% diameter of orbit in males, about 40-45% 
in females; (2) canthal, supraorbital, supratvmpanic, 
postorbital, preorbital, pretympanic, and supralabial 
crests present; (3) cranial crests well developed, robust, 
except parietal crest thin, low, or absent; (4) tibia short, 
about 34% SVL; (5) feet short, about 34 % SVL; (6) dorsal 
tubercles medium to large-sized, elevated, rounded, 
densely arranged on middorsal region, becoming large 



12 



SciENTiiic Pai'i:rs. Naturai History Mi siim.The UNiviRsin oi Kansas 



and conspicuously spinosc Litciallv; (7) ventral tubercles 
reiati\el\' large, granular, with conspicuous spinose 
apices; (8) lateral descending row of enlarged tubercles 
absent; (9) skin texture not sexually dimorphic; (10) 
vocal slit unilateral in male; (11) iii. intcrln/oidciifi poorly 
differentiated from )/;. intcniiaiidibuhnis, but differentiated 
posteric^rly forming a large, unilobed vocal sac with 
heavy black pigmentation; (12) snout shape rounded in 
lateral view, pointed in dorsal view; (13) parotoid glands 
round to ovoid and large, about 1.5-2 times size of eyelid; 
(14) skin between cranial crests on top of head with few 
to many scattered, low, rounded tubercles; (15) ventral 
cciloration whitish cream, with scattereci dark brown, 
indistinct to distinct, punctuations in some individuals. 

Biifo ci/tindcii may be distinguished from all other 
members of the Bnfo coccifcr Group by the combination of: 
its relatively small size; relatively large, scattered, rounded 
tubercles on the middorsum; large, spinose tubercles on 
the dorsolateral and lateral surfaces; relatively large, 
spinose tubercles on the venter; large parotoid glands; 
weakly developed (or absent) parietal crest; and presence 
of scattered, dark brown punctations on the venter (but 
this last character is variable amtmg specimens). 

Biifo ci/cladcn differs from B. coccifer by being smaller 
(males of B. coccifcr to 62 mm SVL, females to 82 mm), 
and possessing the following characters: large, elevated, 
densely arranged middorsal tubercles (middorsal tuber- 
cles relatively small, scattered in B. coccifcr); large, 
strongly spinose dorsolateral and lateral tubercles 
(dorsolateral and lateral tubercles smaller in B. coccifcr, 
variably roundeti or moderately spinose); relatively large, 
strongly spinose ventral tubercles (ventral tubercles small, 
rounded, or \sith tiny spinose apices in B. coccifcr); weakly 
de\'eloped, or no, parietal crests (parietal crest relativeK' 
well developed in most specimens of B. coccifcr); and an 
adverti.sement call with a higher pulse rate (Table 1, Fig. 
2). Biifo cycladcn differs from B. pisinnus by having: much 
larger, distinctly spinose dt)rsolateral tubercles (smallei-, 
more rounded in B. pisinnus) and relatively larger 
parotoid glands (1-1.5 times si/e of eyelid in B. pisinnus). 
Superficially, B. cyclndcn resembles B. signifcr, but it is 
smaller (males of B. si^nifcr to 64 mm SVL, females to 
77 mm) and has the following features: a whitish-cream 
venter, with or vvitht)ut scattered dark brown punctations 
(venter with bold brown-black marbling in /'. sis^nifcr); 
and smaller and less spinose tubercles overall (but this 
latter character is somewhat variable and subjective). 
Bufo cyclndoi differs from B. ibarrai bv: being smaller 
(males of B. ibarrai to 82 mm SVL, females to 94 mm); 
having males with conspicuously spinose dorsolateral 
and lateral tubercles; having relali\el\' larger, more 
densely arranged middorsal tubercles (all such tubi'rcles 



relatixely small, scattered, rounded in males of B. ibarrai); 
and having rounded middorsal tubercles in females 
(tubercles spinose in females of B. ibarrai). Bufo ci/cladoi 
differs from B. portcri by: having sharply spinose lateral 
tubercles in males (rounded in males of B. portcri); and 
relatively thin supratympanic crests (large, bulbous in B. 
portcri). Morphometric \ariation is summarized in Table 

Distribution and Ecology. — Bufo cycladcn occurs in 
a relati\eh' narrow ele\ational band along the Pacific 
slope of the Sierra Madre del Sur in Cjuerrero and Oaxaca, 
Mexico (Fig. 7). Most specimens ha\e been collected at, 
or near, Agua del Obispo, Guerrero. Althc)ugh ctimmcinly 
cited in the literature, Agua del Obispo does not appear 
on most maps because it is a private hacienda; it is located 
near the 1000-m contour (17°18' N, 99" 28' W) along the 
o\d high\s'a\' between the towns of Tierra Colorada and 
Chilpancingo, Guerrero. One other series (including 
the holotype, UIMNH 57142) is known from Putia de 
Guerrero, Oaxaca; this locality lies at an elevation similar to 
that of Agua del Obispo, Guerrero, about 150 km (airline) 
eastward on the same slope of the Sierra Madre del Sur. 



100° 








1 


V 

k. 20° 


▲ B.cycladen ^~-~~.^_^^*- 

• B. pisinnus ^"^^-—^ 

1 < 300 m 
1 300-900 m 






1 1 900-2100 m 
^^° WM =• 2100 m 
100 




15° 


Kilometers 

100° 







I ij;. 7. Map ol LiMitiMl Mexico with dot localities tor Bufo cudadcii 
iTid K. pii^iiiiiu!^ (see species account lx>lou) indicating specimens 
■x.iTiiined, which reprc-enls all known Imalities. 



Toads oi- thk Blfo coccii i:k Complex 



13 



Presumnblv, the distribution of this species is continuous 
along the middle-elevations of this slope, between these 
two localities. Campbell and Duellman (2000:19) described 
the habitat at Agua del Obispo as "...an area of scattered, 
small pines and brush that appears to be ecotonal between 
tropical deciduous forest, which is found slightly below, 
and pine-oak forest, which is present in the mountains 
above." Field work in this region by JRM and associates 
in 2000 and 2002 reinforced that B. ci/chulcii seems to be 
restricted to this narrow, ecotonal habitat. In this area, we 
encountered: B. Jiuiriinn^ and B. iiuiniioivtis at the lowest 
elevations between Acapulco and near Las Cruces; only 
B. manuorcits in the vicinity of Tierra Colorada (±M00 m); 
B. iimriinif and B. cycladcn near Agua del Obispo; only B. 
l>crplcxii> in tine Zumpango del Rio/Chichihualco area; 
and only B. occidciitalis at high elevations (over 2000 m) 
near Carrizal de Bravo. Da\'is and Dixon (mtiS) reported 
some additional records of Bufo from along this transect, 
but we have not verified the identifications of those 
specimens. The sporadic appearance of B. iiuirinii^ along 
this transect is noteworthv; we also note that Davis and 
Dixon (1965) did not report this species from anywhere in 
the area. 

Tadpole. — The tadpole of B. ci/chnicii is unknown, as 
are aspects of its reproductive biology. Davis and Dixon 
(1965) reported females collected on 22 June " . . .contained 
numerous small eggs." 

Remarks. — The type locality for this taxon is near 
Putla, Oaxaca, Mexico, which is located at about 750 m 
elevation on the Pacific slope of the Sierra Madre del Sur; 
specimens designated as paratypes were collected at the 
type locality and from around Agua del Obispt^, Guerrero, 
Mexico (Lynch and Smith, 1966). However, these authors 
referred all Mexican populations of B. coccifcr to this new 
taxon (thereby including populations of three different 
species of toads, which we have identified as B. coccifci; 
B. cycladcu, and B. yifiiniiis). Furthermore, it is evident 
from their map (Lyncli and Smith, 1966:fig. 2) that their 
comparative samples of "B. coccifcr" from Guatemala, 
Honduras, and Nicaragua may have included specimens 
that we recognize as B. coccifcr, B. iharrai, and B. portcri. 
Owing to this confusion, the diagnosis of B. cycladcn 
presented in the original description has not been widely 
accepted (Porter, 1967; McDiarmid and Foster, 1981; 
Frost, 1985, 2003). We apply the taxon 6. c\/clndeii only to 
those populations on the Pacific slope of the Sierra Madre 
del Sur of Guerrero and Oaxaca, Mexico. It seems likely 
that the confusion associated with 6. cyclnden is the result 
of authors' having inadvertently compared a variety of 
different species in their efforts to distinguish B. coccifcr 
and B. ciicladcii. Mendelson (2001) provided additional 
comments on this problem. 



Bufo iharrai Stuart 
' Figs. 3, 8, 9 

Hufo iharrai Stuart: I S9. Holotypc: UMM7. 108000. Type kicality: 
.AsiTiMdero San Lorenzo |about 12 airline km sli^hth' East of North of 
LiLipa], Jalapa, GuatemaLi, 1725 ni. 

Bufo coccifcr — Cope, 1887 |in part]; Meyer and Wilson, 1971 |in part]; 
Campbell and Vannini, 1989 |in part]; Campbell, 1999 |in part]; McCranie 
and Wilson, 2002 [in part]. 

Hufo iharrai — Lyndn and Fugler, 1965 |in part]: Porter, 1966; I'rost, 
19S5; Camplx'll, 1999; Mendelson, 2001; Frost, 200.1. 

Bufo luicrotif — Sdimidt and Stuart, 1941 |in part, tor reference to 
UMMZ840H3]. 

Diagnosis. — A large species of Bufo (males to 
82.4 mm SVL; females to 94.4 mm SVL), having the 
following combination of characters: (1) tympanum 
e\'ident externally, about 45'7- diameter of orbit in 
males, about 40' r, in females; (2) canthal, supraorbital, 
supratympanic, postorbital, preorbital, pret\-mpanic. 





Fig. 8. Lateral aspects ot the heads ot adult male specimens of Bufo 
iharrai (upper: KU 58413) and B. pi^iiiuiis (lower: UMMZ WED 233723; 
holot\ pe). Scale bar = 1 cm. 



14 



Scientific Papers, Natural History Museum, The University of Kansas 




Fig. y. Dorsjl and vi-ntrdl dspects of representative adult specimens of Bufo ibiinni from Departamento Guatemala, Guatemala (male, left: UTA 
A-25825, SVL = 69.S mm; female, right: UTA A-47572, SVL = 93.3 mm) and B. pi^imnis from Michoaain, Mexico (male, left: UMM2 115353 [WED 
10971], SVL = 43.5 mm; female, right: UMMZ 121578, SVL = 60.4 mm). 



parietal, and supralabial crests present; (3) cranial crests 
well developed, parietal crests low, thin, sharply angled 
medially; supratympanic crest large, bulbous; (4) tibia 
short, about 40% SVL; (5) feet short, abi>ut 40';, SVL; (6) 
middorsal tubercles sparse, roundecl, becoming spinose 
lateralh' in females, all dorsal tubercles in males rounded, 
usually indistinct or absent lateralh'; (7) wntral tubei-cles 
areolate, non-spinose in males and finelv spinose in 
females; (8) lateral descending row of enlarged tubercles 
indistinct or absent; (9) skin textinv sexualK- dimoiphic; 
(10) vocal slit unilateral in male; (11) ni. iiilcr!n/oulru> 
poorly differentiated from ;;;. iiilciiiuiiKlibitliiri^. but 
differentiated posteriorly, forming a large, unilobed vocal 
sac with variable amounts of pigmentation; (12) snout 
shape rounded in lateral \'ievv, weakK' pointed in dorsal 
view; (13) parotoid glands large, ovoitf, length about 
2 times size of eyelid; (14) skin between cranial crests 



usually smooth, lacking tubercles; (15) \'entral coloration 
dull cream with some diffuse gra\' mottling in some 
inctividuals. 

Biifo ibarrai is the largest species in the B. coccifer Croup 
and differs from all species except B. portcri bv: ha\'ing 
i"elati\el\' massi\e cranial crests, with the supratympanic 
crest being distinctb' bulbous; and ha\ing sexually 
dimorphic skin texture with distincth' roimded lateral 
tubercles in males and spinose lateral tubercles in females. 
Bufo ihiiiiiii closely resembles B. porlcvi but differs b\: being 
larger (males to 82.4 mm SVL and females to <-)4.4 mm SVL 
\s. S^.M mm in males and 76.2 mm in females); and ha\ing 
the caudal muscuLiture ot the tadpole uniloini pale blown 
(cream and hea\il\' punctated with brown in B. portcri). 
Morphometric \ariation is summarized in Table 2. 

Distribution and ecology. — Bufo ihnrrni occurs at 
moderate ele\ations (1360-|9S() m) in the highlands of 



Toads of the Blfo coccii kk Compi.bx 



15 




Fig. 10. Map of Guiiteniiila and western Honduras with dot 
localities for Bufo ibnrnii and B. poilcn (see species account below) 
indicating specimens examined. 

central and southern duatemala (Mendelson, 2001), and 
new information presented here extends that range into 
contiguous regions of Honduras (Fig. 10). The Honduran 
localities range up to 2020m elevation, and all represent 
Premontane Moist Forest (e.g., McCranie and Wilson, 2001: 
pl.2E) or Lower Montane Moist Forest (e.g., McCranie 
and Wilson, 2001:pl.4E) and lie in the western ranges of 
the Southern Cordillera Region (sensu McCranie and 
Wilson, 2001), in the departments of Intibuca, Lempira, and 
Ocotepeque (Fig. 10). These discoveries refute Mendelson's 
(2001 ) premature speculation that B. ibnvrni ma\' be endemic 
to Guatemala and refute McCranie and Wilson's (2001) 
conclusion that B. ibanai does not occur in Honduras. 

Tadpoles. — The tadpole was described bv Mendelson 
(2001 ). 

Remarks. — Altliough Mendelson (2001) discussed 
diagnostic features that distinguish B. coccifcr and B. iharrai, 
he did not present multi\ariate analyses of morphometrv. 
In addition, Mendelson's (2001) research was based only 



on specimens from Guatemala. During the course of this 
stud\', Eric N. Smith kindly sent to us specimens of B. 
iharrai he collected in Ocotepeque, Honduras. These toads 
are consistent with the diagnosis presented b\ Mendelson 
(200 1 ) and have a similar (0.67' <'. di\-ergent) mitochondrial- 
DNA sequence to B. iharrai collected at the opposite end of 
the range of the species, in El Quiche, Guatemala. We also 
discovered additional specimens from western Honduras 
among museum collections (Appendix II) that match tiie 
diagnosis provided b\' Mendelson (2001 ); these specimens 

Bufo pisinuus new species 
Figs. 3, 8, 9 

Bufo coccifcr Porter, 1%3 [in part]; Frost, 1985 |in part]; Villa et al., 
mSS |in part]: Flores-Villela, IW.I |in part]: Campbell, 1999 |in part]; 
I i-ost, 2003 |in part], 

Bufo ci/clihicn Lvnch and Smith, l"-l(i(i |in part, tor reference to 
specimens from Michoacan, Mexico]. 

Holotype.— UMMZ 233723 (WED 10973), an adult 
male from 6.2 mi [10 km] E Apatzingan, 1100 feet [335 m] 
elevation, obtained b\' W. E. Duellman and R. E. Etheridge 
on 2 August 1936. 

Paratypes. — All from Michoacan, Mexico: 6.2 mi [10 
km] E Apatzingan, 1100 ft [333 m] (UMMZ 115353 [15 
specimens]); 7 mi [11.2 km[ E Apatzingan, 1100 ft [335 m] 
(UMMZ 112794 [6 specimens), 115335); 1 mi [1.6 km[ W 
Apatzingan, 1100 ft [335 m] (UMMZ 115354); 3 mi [4.8 
km] S Lombardia (UMMZ 121578). 

Diagnosis. — A small species of Bufo (males to 51 
mm SVL; females to 62 mm SVL), having the following 
combination of characters: (1) tympanum evident , 
about 35^0 '}o diameter of orbit in males, about 40-50% 
in females; (2) canthal, supraorbital, supratympanic, 
postorbital, preorbital, pretympanic, supralabial crests 
present, parietal crest a thin ridge, or absent; (3) cranial 
crests weakly developed, thin, low; (4) tibia short, about 
34';;, SVL; (5) feet short, about 36% SVL; (6) dorsal 
tubercles small to medium sized, low, rounded, relatively 
densely arranged middorsally, becoming smaller and 
spinose laterally; (7) ventral tubercles tiny, evenly 
granular, appearing smooth, especially in males, but tiny, 
spinose apices apparent under microscope; (8) lateral 
descending row of enlarged tubercles absent; (9) skin 
texture not sexually dimorphic; (10) vocal slit unilateral 
in male; (11) ;;;. iiitcrln/oideus poorly differentiated from ;;/. 
iutcniiaiidihiilaris, but differentiated posteriorly forming a 
large, unilobed vocal sac with heavy black pigmentation; 
(12) snout shape acutely rounded in lateral view, sharply 
pointed in dorsal view, snout shape in females similar but 
more genth' rounded in lateral view; (13) parotoid glands 
ovoid, about 1.0-1.5 times size of eyelid; (14) skin between 
cranial crests on top of head usually with many scattered, 
low, rounded tubercles; (15) ventral coloration usually 



16 



Scii-Miiic Paplrs, Nailkal History Museum, Thh Umvl;rsha oi Kansas 



immaculate whitish cream, some indixiduals with tinv, 
scattered black flecks. 

Biifo pnsiiiiitif can be distinguished from all other 
members of the B. coccifcr Group b\' its small si/e, and 
possession of: relatively weakly developed cranial crests, 
the parietal crest appearing only as a thin ridge among 
surrounding tubercles and frequentlv is absent; smaller, 
more densely arranged dorsal tubercles; and smaller, 
less spinose ventral tubercles. Bitfo pisinniis differs from 
B. coccifcr by having: a relatively shorter snout and an 
advertisement call with a higher pulse rate (120 pulses 
per sec vs. maximum of 115 pulses; Table 1, Fig. 2). 
Biifo pisi)uni^ differs from B. cycladcn by having: much 
smaller, and less spinose dorsolateral tubercles (large 
and conspicuously spinose in B. cycladcn); and smaller 
parotoid glands (about 1.5-2.0 times size of eyelid in B. 
cyclndcn). Biifo p'ifiiiniis differs from B. sigiiifcr by having: 
a whitish-cream venter, with or without scattered black 
flecks (venter with bold brown-black marbling in B. 
sigiiifcr); smaller parotoid glands (about twice size of 
eyelid in B. sigiiifer); and an advertisement call with a 
higher frequency and faster pulse rate (Table I, Fig. 2). 
Bufo pisiimiis differs from B. ibarrai by having: smaller, 
more rounded parotoid glands (glands in B. ibnrnii ovoid, 
much higher, and about twice size of the eyelid); and 
rounded mid-dorsal tubercles in both sexes (tubercles 
in B. ibarrai rounded in males, spinose in females). Bufo 
pisiunus differs from B. portcri by having: sharply spinose 
lateral tubercles in males (rounded in males of B. portcri); 
and a relatively thin supratympanic crest (large, bulbous 
in 6. portcri). 

Description of holotype. — Body robust; head wider 
than long, width 3M.1';, SVL, length 35.3";- SVL; snout 
sharply pointed in dorsal view, rounded in profile, 
rostral keel distinct; canthal, preorbital, supraorbital, 
pretympanic, supratvmpanic, and postorbital crests 
present, distinct; parietal crests reduced, barely distinct; 
skin on top of head co-ossified; nostril not protuberant, 
directed dorsallv; can thus rostralis forming distinct, raised, 
canthal crest; loival region cinna\'i-; lip distinct, rounded; 
suborbital crest present, distinct, extending from angle of 
the jaw anteriorlv to level of anterior margin ot orbit; notch 
at symphysis of upper jaw pivsenl, distinct; e\'e-nostril 
distance 5. 7'.r, diameterof orbit; t\-mpaniim distinct, nearh' 
round; tympanic annulus distinct onl\' along anterior and 
ventral margins, upper margin contacting supratympanic 
crest, posterit)r margin obsciu-ed by overlying llesh. 
Forelimb short, robust; hand broad, with short, slender 
fingers; relative length ot fingers: II • I < IV ■ III, webbing 
and lateral fringe on fingers absent; tips ot lingers not 
enlarged, smooth dorsallv, demarcated proximalK' b\- 
distinct dermal told; palmar tubercle elistincl, large, 
ovoid; pollical tubercle smaller than palmar tubercle. 



ovoid; subarticular tubercles distinct, elevated, triangular 
in profile, single except distal tubercle on Finger 111 bilid; 
supernumerary tubercles of unequal size, small, distinct, 
scattered evenly over palm and ventral surfaces of 
fingers; nuptial excrescences present as brown granular 
patches on medial surfaces of Fingers I and II. I iind limbs 
short, slender, tibia length 35.8',' SVL; foot length 38.0% 
SVL; tarsal fold absent; outer metatarsal tubercle minute, 
elevated, ovoid; inner metatarsal tubercle slightly larger 
than outer metatarsal tubercle, distinctly elevated, ovoid; 
toes long, slender, relative lengths of toes: I < 11 < V < III 
< IV; lateral fringe present on Toes II, 111, and \', absent 
on Toes I and IV; webbing thin, webbing formula 11 — 
3II2— 3III2— 4IV4— 2V; tips of toes not enlarged, smooth 
dorsally, demarcated proximally by distinct dermal fold; 
subarticular tubercles distinct, elevated, triangular in 
profile, bifid; supernumerar\' tubercles unequal in size, 
distinct, distributed evenlv over \-entral surfaces of foot 
and toes. 

Skin on dorsum of bodv rugose with e\enl\' 
distributed, small, rounded tubercles of relatively equal 
size, becoming sharpiv pointed laterally; parotoid glands 
about same size as evelids, o\oid, oriented perpendicular 
to midline of bodv; lateral rov\' of enlarged tubercles 
barely evident; dorsal surface of head smooth with few, 
small, rounded tubercles scattered in interspaces between 
cranial crests; dorsal surfaces of limbs covered with small, 
weakly pointed tubercles; skin on throat and other ventral 
surfaces granular, covered with tinv flattened and weakly 
pointed tubercles. 

Choanae small, rounded, widely spaced; teeth and 
odontoids absent; tongue long, ovoid, about four times 
as long as wide, free posteriorly for about one-fourth its 
length; vocal slit unilateral, sinistral. 

Coloration of holotype. — In preser\ati\e (ethanol), 
dorsum ot bod\' and limbs mottled evenly with pale brown 
and dark brown markings; irregular cream blotches present 
posterior to each parotoid gland; top ot head uniform dark 
brown with distinct cream interorbital bar; distinct cream 
middorsal stripe extending from snout to posterior end 
o\ urostvie, irregularlv widened at se\'eral points along 
its length; lateral sin-faces pale brown with dark brown 
flecking. Ventral surfaces nearh' immaculate cream: dark 
\'ocal sac visible through gular skin. 

Measurements of the holotype (in mm). — SVL 45.0, 
HI. IS», IIW 17.(1, IL Ih.l. 14 17.1, orbit diameter 5.4, 
tympanum diameter 23, suprahmpanic crest length 2.7, 
parotoid gland length ^.4, pai'otoid gland width 4.8. 

Coloration in life. — Duellman (l'-)6l:2I) described 
coloration "...yellowish tan ground color with dark brow n 
spots; middorsal stripe is deep ncIIow or ci-eam color. 1 he 
\ enter is dust\- cream color, anil the iiis is pale gold." 



Toads of the Bcro cocciier Complex 



17 



Variation. — Morphometric Vciriation among specimens 
examined is summarized in Table 2. The parietal crest is 
poorly developed in most specimens, and essentially 
absent in a few individuals (e.g., UMMZ 121578). Some 
individuals (e.g., UMMZ 112794) have a dark-brown 
ground color on the dorsum, which effectively obscures 
the dark brown dorsal blotches; most specimens have a 
pale brown to grayish ground color on the dorsum, and the 
overlying brcnvn blotches are distinct. The brown blotches 
on the dorsum may be relativelv small, incorporating one 
to three dorsal ttibercles (e.g., UMMZ 115353 [WED 10972]), 
or the blotches may be larger, incorporating more than 12 
dorsal tubercles (e.g., UMMZ 115353 [WED 10974]). The 
narrow middorsal cream stripe invariably is present in all 
individuals examined. Scattered black flecks on the ventral 
surface may be absent {c.;^., UMMZ 121578), restricted to 
the peripheral ventral surfaces (e.g., UMMZ 1 15353 [WED 
10970]), or scattered relatively evenly across the venter (e.g., 
UMMZ 115354 [WED 10976]). 

Etymology. — The name pisiiiiiiis is the Latin word 
pi>iuiiii<, meaning small, in reference to the diminuti\'e 
size of this species. 

Distribution and ecology. — Bufo pisiiuiii^ is known 
only from the Tepalcatepec Valley, which represents the 
western extension of the Balsas Basin (Fig. 7). Duellman 
(Field Notes, 16 July 1960) described the area arciund 
Lombardia, Michoacan, as having grassy areas interrupted 
by areas of mescjuite with many reddish rocks and barren 
areas. Duellman (1961) suggested that this species is likely 
widespread in region, but noted that it does not occur along 
the coast of Michoacan. Duellman ( 1 961 ) reported breeding 
choruses in muddy ditches and flooded grassy fields after 
heavy rains in June and August. 

Tadpoles. — The tadpole of B. pifiiniiiis is unknown. 

Bufo porteri new species 
Figs. 3, n, 12 

Bufocoi'iifcr — MevtTiind Wilson, l'-)7I |in p>irt|; iTost, 198S [in portl; 
McCranie dnd WiKon, 2002 |in part): Campbull, Iwm |in part]: Frcist, 20(1^ 
|in part|. 

Bufo ibarnii — l.vnch iind Fiigler, I'-Ui? |in part]. 

Holotype. — KU 97519, and adult male from 6 mi 
[9.6 km] NE Escuela Panamericana, Cerro Uvuca, 520(1 ft 
[1584 m], Francisco Morazan, Honduras, obtained by K. 
R. Porter on 30 June 1964. 

Paratypes. — All from Franciso Morazan, Hondinas: 
6 mi ]9.h km] NE Escuela Panamericana, Cerro Uvuca, 
5200 ft [1584 m] (KU 97520-26); 6 mi 19.6 km] NE Escuela 
Panamericana, Cerro Uvuca, 6000 ft 11828 m] (KU 97514); 
W slope Cerro Uyuca, 1650 m (KU 103221); Parque 
Nacional LaTigra aLxiveSan Juancito, 2100 m (KU 192294); 
Cerro La Tigra, 1840-1890 m (KU 194216-19, 209249). 








Fig. 11. Lateral aspects of the heads of adult male specimens of 
Bufo portcn (upper; KU 97520) and B. ngnifcr (lower: AMNH 69626). 
Scale bar = 1 cm. 

Diagnosis. — A small to medium-sized species of 
Blip (males to 59.9 mm SVL; females to 76.2 mm SVL), 
having the following combination of characters: (1) 
tympanum evident externallv, about 40',, diameter of 
orbit in both males and females; (2) canthal, supraorbital, 
suprat\'mpanic, postorbital, preorbital, pret\'mpanic, 
parietal, and supralabial crests present; (3) cranial crests 
well developed, parietal crests low, thin, supratympanic 
crest large, bulbous; (4) tibia short, about 37' ,'. SVL; 5) feet 
short, about 37"^. SVL; (6) middorsal tubercles sparsely 
arranged, rounded, becoming spinose laterallv in females, 
all dorsal tubercles in males rounded, usually bect)ming 
indistinct or absent laterallv; (7) ventral tubercles areolate, 
non-spinose in males, fineh' spinose in females; (8) lateral 
descending row of enlarged tubercles indistinct or absent; 
(9) skin textu re sexuall\'dimorphic;( 10 )\ocal slit unilateral 
in male; (11) iii. iiitciin/ouicu> poorly differentiated from 
III. iiitcniiiiiidibiilnrifi differentiated posteriorly, forming a 



Scientific Paphrs, Naturai. Hisiory Museum. The University of Kansas 




Fig. 12. norsjj iind ventral aspects ot representative adult specimens of Biifo portcrl from Francisco Morazan, Honduras (male, left: KL' ^7519, 
SVL = 51.6 mm; female, right: KU y7.S14, SVL = 60.7 mm) and B. signifcr Panama (adult male, left: TNHC 31341, SVl, = 49.7 mm; subadult female, 
right: KU 1 1 3359, SVI . = 54. 1 mm ). 



Itirgc, unilobed vocal sac hcavih' pii^LMiiented in black; 
(12) snout shape roiindccl in lateral \ic\v, wcakh' poifited 
in dorsal xiew; (13) pafotoid glatids moderate, rounci to 
ovoid, length about 1.00-1.23 titnes size of eyelid; (14)skifi 
between cranial crests usually smooth, lacking tubercles; 
(15) ventral coloration is dull cream with some diffuse 
gray mottling in some individuals. 

Biifo poiicii ma\' be distingiushed trotn all other 
tnember of file B. coaifcr Group, except B. ihninu, the 
sexual dimorphism in the texture of the dorsal skin 
and the relatively robust cratiial crests, especialh' the 
supratympanic crest. Biifo poiicri closely resetnbles />'. 
ibanni but differs by being smaller (tnales to 54.9 mm 
SVL and females to 76.2 mm SVL \'s. males to >S2.4 mm 
SVL and females to 94.4 mtii SVL); adult male B. ]wrtcn as 
small as 43.9 mm SVL have been observed (e.g., LSUMZ 



46431 ). The caudal musculature of the tadpole of B. poiicri 
is creafTi w ith hea\ \ brown punctations (See description 
b\' McCranie and Wilson, 2002:173; caudal musculature 
if! B. ibarrai is unifortii pale browti.) 

Description o( holotype. — Bod\ robust; head \\ ider 
than long, width 40.0% SVL, length 36.4% SVL; snout 
sharply pointed in dorsal view, pointed in profile, rostral 
keel distinct; canthal, preorbital, supraorbital, pretytnpanic, 
supratympatiic, and postorbital crests present, distinct; 
supraorbital atid suprat\'mpanic crests distintly thickened; 
parietal crests present, not reduced; skin on top of head 
co-ossified; nostril protuberant, directed dorsally; canthus 
rostralis forming distinct, raised, canthal crest; loreal region 
concaM'; lip distinct, rounded; suborbital crest present, 
distinct, extending from angle of the jaw anteriorly to le\'el 
of afiterior margin of orbit; notch at s\'mphvsis of upper 



Toads of ihh Buio coven ik Complex 



19 



jaw present, distinct; e\'e-n(istril distance 37.9",- diameter of 
orbit; tympanum distinct, nearly round; tympanic annulus 
distinct only along anterior, xentral, and posteroventral 
margins; upper margin of tympanic annulus contacting 
and obscured by supratvmpanic crest. Forelimb short, 
robust; hand broad with short, slender fingers; relative 
length of fingers: II < I < IV < III, webbing and lateral 
fringe on fingers absent; tips of fingers not enlarged, 
smooth dorsallv, demarcated proximally by distinct 
dermal fold; palmar tubercle distinct, large, rounded; 
pollical tubercle smaller than palmar tubercle, rounded; 
subarticular tubercles distinct, ele\ated, triangular in 
profile, all single except distal tubercle on Finger 111 bifid; 
supernumerary tubercles of unequal size, small, distinct, 
scattered evenly over palm and ventral surfaces of fingers; 
nuptial excrescences present as brown granular patches 
on medial surfaces of Fingers l-IIl. Hind limbs relatively 
long, slender, tibia length 41.7",. SVL; foot length 40.5",'. 
SVL; tarsal fold absent; outer metatarsal tubercle very 
small, rouncied, indistinct; inner metatarsal tubercle much 
larger than outer metatarsal tubercle, distinctly elevated, 
ovoid; toes long, slender, relative lengths of toes: I < II < 
V < III < IV; lateral fringe barely evident on Toes III and 
IV, absent on other toes; webbing thin, webbing formula 
II — 3II2 — 3III2 — 4IV4 — 2V;'tips of toes not'enlarged, 
smooth dorsally, demarcated proximally bv distinct dermal 
fold; subarticular tubercles distinct, elevated, triangular in 
profile, bifid on Toes III and IV; supernumerary tubercles 
unec^ual in size, distinct, distributed evenly over ventral 
surfaces of foot and toes. 

Skin on dorsum of body relatively smooth with 
scattered, small, rounded tubercles of unequal size, many 
bearing tiny, indistinct single keratinized apices; tubercles 
on lateral surfaces indistinct, roimded or ovoid, ncit 
pciinted; parotoid glancis about same size as eyelids, ovoid, 
oriented slightly divergent tci midline of body; distinct 
lateral row of enlarged tubercles absent; dorsal surface of 
head smooth \yitlT few, small, rounded tubercles scattered 
in interspaces between cranial crests; dorsal surfaces of 
limbs covered with small weakly pointed tubercles; skin 
on throat and other ventral surfaces smooth!)' granular. 

Choanae moderate in size, rounded, widely spaced; 
teeth and odontoids absent; tongue long, ovoid, about four 
times as long as wide, free posteriorly for about one-foiuih 
its length; vocal slit unilateral, dextral. 

Coloration of holotype. — In preservative (ethanol), 
bod\' pale brown with indistinct, medium-brown marbled 
markings diffused across middorsal and lateral areas; 
distinct, thin cream middorsal stripe present; two oblong 
dull cream patches on dorsolateral areas; distinct dark 
brown bar extending across area between parietal crests 
and another covering area between canthal crests; all 
limbs with indistinct medium brown crossbars; venter 



immaculate dull cream; dark vocal sac clearly \'isible 
through gular skin. 

Measurements of the holotype (in mm). — SVL 51.3, 
HL 18.7, HW 20.5, TL 21.4, FL 20.8, orbit diameter 7.6, 
tympanum diameter 3.0, supratympanic crest length 3.1, 
parotoid gland length 6.9, parotoid gland width 4.8. 

Coloration in life. — McCranie and Wilson (2002:pl 
'■'D) provided a color photograph of this speceis; see also 
Figure 3. 

Variation. — Morphometric variation among specimens 
examined is summarized in Table 2. The parietal crests 
in females generally are well developed and distinct; 
however, the condition of the crest varies among males 
and may be relatively robust (e.g., KU 209249) or reduced 
to an indistinct, thin sliver of raised bone (e.g., KU 97523). 
The brown dorsal blotches may be very dark brown, and 
therefore quite distinct (e.g., LSUMZ 46398), or they may 
be only slightly darker than the brown ground cc^lor of 
the dorsum (e.g., LSUMZ 46445). The ventral coloration 
is either uniform dull cream (e.g., KU 103221) or bearing 
diffuse, grayish-brown marbling that ranges from 
moderate (e.g., KU 97522) to extensive (e.g., KU 194216). 
The middorsal cream stripe invariably is present on all 
individuals examined; this stripe is quite indistinct in a 
small nimiber of individuals (e.g., KU 97520) and may 
appear irregular (i.e., not ft)rming a straight line; e.g., 
LSUMZ 45441). 

Etymology. — The specific epithet is a patronym that 
honors Kenneth R. Porter and his series of papers (e.g.. 
Porter, 1963, 1964, 1965)onthesystematicsof Mesoamerican 
Biifo, and also recognizes his numerous field efforts that 
resulted in many of specimens referred to herein. 

Distribution and ecology. — Bitfo povtcvi is known 
from the Hondiu-an departments of Comavagua, Francisco 
Morazan, and La Paz (Fig. 10). The known localities for this 
species generally represent Lower Montane Moist Forest 
habitats (e.g., McCranie and Wilson, 2001 :pl. 4C) in the 
Montanas de Comayagua region. These localities differ 
markedly from the lower elevation, Dr\' and Arid Forest 
habitats (e.g., McCranie and Wilson, 2001 :pl.lC) occupied 
b\' B. coccifer. 

Tadpoles. — Tadpoles leferrable to B. yoiicri (based 
on geography) were described bv McCranie and Wilson 
(2002:173). The tadpole of this species resembles that of B. 
coccifer (McDiarmid and Foster, 1981; Sa\age, 2002), from 
which it is distinguished by having submarginal papillae 
on the oral disc (absent in 6. coccifer) and cream-colored 
caudal musculature with brown punctations (boldly 
marked with brown saddles in B. coccifer). 

Remarks. — We have allocated many Honduran 
specimens that previously were referred to B. coccifer 



20 



Scientific Paphrs, NATiiRAi. Hisiok^ Mu.shum,Thh UNivi-.Rsrn oi Kansas 



(e.g., McCranie and Wilson, 2001) to the new taxon B. 
porteri, or to B. ibarrai (discussed above, and Appendix II). 
The difficulty of identifying specimens from this country 
is exacerbated bv the resemblance of B. porteri and B. 
ibarrai. The wide range of variation among specimens of 
"B. coLxifcr" from Honduras described by McCranie and 
Wilson (2001 ) seems to be attributable to the fact that three 
relative!)' similar species occur in close proximit)' in that 
country. 

At a general le\el, this species appears to be 
parapatric with respect to the distribution of Bnfo coccifcr. 
Thus, the distribution and habitat associations of this 
species, with respect to those of B. coccifer, resemble the 
relationship between B. coccifcr and B. ibarrai in Guatemala 
as described by Mendelson (2001 ). Inasmuch as species 
of Bnfo frec]uentl\- are interfertile (Blair, 1963; Masta et al., 
2002), it is possible that B. porteri may hybridize with B. 
coccifer if the species co-occur on the lower slopes of the 
Pacific Versant of Honduras. Similarly, hvbrids between B. 
porteri and 6. ibarrai eventually may be found. 

Bnfo sigtiifer new species 
Figs. 3, 11, 12 

Bufo coccifer Dunn, 1933; Zweifel, 1965; Frost, 1985 [in part, tor 
reference to specimens from Panama]; Villa et al., 1988 |in part, for 
reference to records from Panama]; Campbell, 1999 |in part]. 

Holotype.— AMNH 69626, an adult male from 7 mi N 
[11.2 km] and 2 mi [3.2 km] W of David, Chiriqui', Panama, 
obtained bv R. G. Zweifel on 25 June 1962. 

Paratypes. — All from Panama. Chiriquf: 7 mi [ 1 1 .2 km[ 
E ConcepcicSn (AMNH 69627); 2.5 mi [4.0 km| NE David 
(TNHC 3134-43); 23 km NNE San Felix, 900 m (USNM 
297511-21). Code: El Valle de Anton (AMNH 59634, 59637); 
16 km S, 9 km W Penonome, 30 m (KU 11 5359-6 1 ); 3.2 km 
W Agua Dulce, 15 m (KU 115362). Herrera: Jacinto, 2250 ft 
[686 m] (ANSP 22341-44); 3 mi [4.8 km] SW Pan American 
Hwy, on road past Potuga (UMMZ 167373). Veraguas: 14 
km NE Sona, 75 m (KU 95432). 

Diagnosis. — A medium-sized species of Bufo 
(males to 64 mm SVL; females to 77 mm SVL), ha\ ing 
the following combination of characters: (1) tympanum 
evident externally, about 40-45'('. diameter of orbit in 
males, about 40-50% in females; (2) canthal, supraorbital, 
supratympanic, postorbital, preorbital, pret\nipanic, 
parietal and supralabial crests present; (3) most cranial 
crests distinct and thick, except parietal crests low, thin, 
sometimes intermittent; (4) tibia short, about 35'f' SVL; (5) 
feet short, about 36' V, SVF; (6) dorsal tubercles medium- 
sized, prominent, roimded, rel,ili\el\' denseh' arranged 
middorsally, becoming smaller, more concentrated, antl 
spinose laterally; (7) ventral tubercles granular, willi 
small, distinct spinose apices; (8) lateral desci'nding row 
of enlargeti tubercles absent; (9) skin texture not sexualh' 



dimorphic; (10) vocal slit unilateral in male; (11) iii. 
iiiterh\/oiiieiis poorly differentiated from iii. iiiteriiniiulibulari^, 
but differentiated posteriori)' forming a large, unilobed 
vocal sac with heavy black pigmentation; (12) snout 
shape rounded in lateral view, pointecf in dorsal \'iew; 
(13) parotoid glands round to subovoid, about twice size 
of e\'elid; (14) skin between cranial crests on top of head 
usually with few, scattered, low, rouncfed tubercles; (15) 
ventral coloration cream with distinct, bold, brown-black 
marbled pattern, becoming indistinct o\er pehic patch. 

Bnfo ^i^'^iiifer may be distinguished from all other 
members of the Bufo coccifer Group bv ha\'ing a cream 
venter o\'erlain with a distinct, marbled pattern of brown- 
black markings. All t)ther species in the group have 
immaculate, ornearh' immaculate, cream ventral surfaces 
with the exception that some indi\iduals of B. cucladeu may 
ha\'e some dark brown mottling, and some indix'iduals of 
B. pisinniis mav ha\'e some tiny, black flecks. Bufo si^i^uifer 
differs from B. coccifer by having: relatixel)' thinner and 
lower parietal crests (typically higher and thicker in B. 
coccifer); relatively smaller tympana; and an advertisement 
call with a lower frequencv and pulse rate (Table 1, Fig. 2). 
Bufo :^igiiifer is superficially quite similar to B. ci/cladeu but 
differs by: being larger (males of B. cycladen to 54 mm SVL, 
females to 62 mm); having a distinctly marbleci \'entral 
pattern (variably present, but always less developed in 
B. cycladen); and having generallv smaller, less spinose 
tubercles overall (but this latter character is someu'hat 
\'ariable and subjective). Bnfo signifer differs from B. 
pisinnub bybeing larger (males of B. pi^inuns to 51 mm 
SVL, females to 62 mm) and by having: larger middorsal 
tubercles (small in B. pisinnns); larger parotoid glands 
(glands about 1.0-1.5 times size of eyelid in B. pisiniius); 
spinose tubercles on the venter (granular, non-spinose in 
B. pisi)i}uis); an advertisement call with a lower frequency 
and pulse rate (Tablel, Fig. 2); and better-developed cranial 
crests, especially with respect to the parietal crest (crests 
weaklv developed, and parietal crest very reduced or 
absent in B. pissiiin^). Bnfo signifer differs from B. ibarrai by 
being smaller (males of B. ibarrai to 82 mm SVL, females 
to '^'4 mm) and b\' having: smaller, more rounded parotoid 
glands (glands in B. ibarrai ovoid, about twice the size of 
the e\'elid); and rounded mid-dorsal tubercles in both 
sexes (tubercles in B. ibarrai roimded in males, spinose 
in females). Bnfo fiignifer differs trom B. porteri by having: 
sharplv spinose lateral tubercles in males (roimded in 
males of B. porteri); and a relatively thin supratympanic 
crest (large, bulbous in B. porteri). 

Description of holotype. — Body robust; head wider 
than long, width 39.9', SVL, length 34.7';;, SVL; snout 
sliarph' pointed in dorsal \'iew, rounded in profile, rostral 
keel distinct; canthal, pivorbital, supraorbital, pretympanic, 
siiprat\mpanic, and postorbital crests present, distinct; 



Toads of the Bufo coccih.r Complex 



21 



parietal crests present, relati\el\' indistinct; skin on top 
of head coossified; nostril protuberant, directed dorsally; 
canthus rostralis forming distinct, raised, canthal crest; 
loreal region concave; lip distinct, rounded; suborbital 
crest present, distinct, extending from angle of the jaw 
anteriorly to level midway between orbit and nostril; notch 
at symphysis of upper jaw present, shallow, indistinct; 
eye-nostril ciistance 47.6' r. diameter of orbit; t\mpanum 
distinct, nearlv round; t\mpanic annulus distinct only 
along anteroventral margin, upper margin contacting 
supratympanic crest, posterior margin obscureci by 
overlying flesh. Forelimb short, robust; hand broad, with 
short, slender fingers; relati\'e length of fingers: II < I < 
IV < III, webbing and lateral fringe on fingers absent; 
tips oi fingers not enlarged, smooth dorsallv, demarcated 
proximalh' by distinct dermal fold; palmar tubercle 
distinct, large, round; pollical tubercle smaller than palmar 
tubercle, ovoid; subarticular tubercles distinct, elevated, 
triangular in profile, single except ciistal tubercle on Finger 
III bifid; supernumerary tubercles of unequal size, large, 
distinct, scattered evenly over palm and \entral surfaces 
of fingers; nuptial excrescences present as bro\N'n granular 
areas on entire dcirsal surfaces of Fingers I and II, including 
lateral surfaces of tips of fingers, lateral surfaces of distal 
phalange of Finger III, and on medial surface of pollical 
tubercle. Hind limbs short, slender, tibia length 34.5' «'. SVL; 
foot length 35.1"o SVL; tarsal fold absent; outer metatarsal 
tubercle small, elevated, ovoid; inner metatarsal tubercle 
slightly larger than outer metatarsal tubercle, distincth 
elevated, ovoid; toes long, slender, relative lengths of toes: 
I < II < V < III < IV; lateral fringe on toes barely evident 
on Toes II and III, absent on other toes; webbing thin, 
webbing formula II — 27,111'/, — 3III2 — 4IV4 — 2V; 
tips of toes not enlarged, smooth dorsally, demarcated 
proximally by distinct dermal fold; subarticular tubercles 
distinct, elevated, triangular in profile, single except two 
most distaltubercles on Finger IV and most distal tubercle 
on Finger V bifid; supernumerary tubercles unequal in 
size, distinct, distributed evenly over ventral surfaces of 
foot and toes. 

Skin on dorsum of body rugose with scattered, 
rounded tubercles of unequal size, becoming sharply 
pointed laterally; parotoid glands about same size as 
eyelids, slightly ovoid, oriented parallel to midline of 
body; lateral row of enlarged tubercles barely evident; 
dorsal surface of head smooth with few, small, rounded 
tubercles scattereti in interspaces between cranial crests; 
dorsal surfaces of limbs covered with small to large, mostly 
pointed tubercles; all ventral surfaces rough, co\'ered with 
small, conical tubercles. 

Choanae small, ovoici, widely spaced; teeth and 
odontoids absent, but ventral surface of neopalatine 
appears serrate; tongue long, ovoid, about four times 



as long as wide, free posteriorly for about one-fourth its 
length; \'ocal slit unilateral, dextral. 

Coloration of holotype. — In preservative (ethanol), 
dorsal areas of body and limbs mottled evenly with pale 
brown and dark brown markings, with a bilateral series of 
small, oblong cream markings dorsolaterally; top of head 
uniform dark brown with distinct cream interorbital bar; 
distinct cream middorsal stripe extending from snout to 
posterior end of urostyle; lateral surfaces dull brown with 
some dull cream supralabial spots. Ventral surfaces dull 
cream with indistinct pale gray markings, forming loosely 
reticulate pattern; dark vocal sac barely visible through 
gular skin. 

Measurements of the holotype (in mm). — SVL 57.1, 
HL 20.5, HVV 23.0, TL l'-).9, FL 20.0, orbit diameter 8.2, 
tympanum diameter 4.4, supratympanic crest length 3.4, 
parotoid gland length 6.8, parotoid gland width 5.9. 

Coloration in life. — Original field notes by R. G. 
Zweifel { 1 2 June 1 962) describe AMNH 69625 as " . . .a rather 
black and white looking toad, the whiteness coming from 



80° 



10° 




B. agnifer 



< 300 m 
300-900 m 
900-2100 m 



I ""H > 2100 m 
100 

Kilometers 



80= 



Fig. 13. Map of western Panama with dot localities for Biifo >iguiffr 
indicating specimens examined, which represent all known localities. 



97 



SciHNTiHC Paphrs. Natural History Museum, The University of Kansas 



the lateral stripe which begins at tlie parotid and runs back 
to the groin and by the central vertebral line. The blackness 
is supplied by the dorsal blotches which are almost jet- 
black. The ground color between the blotches is dark gray. 
The side below the lateral line is a mixtiuv ot black and 
gray. The eyelids are a stripe with grayish white and black. 
The interorbital area is black with a grayish white band. 
The \'entral surfaces are mottleci gray and white; there is 
a dark gray spot in the middle posterior gular region. The 
hind legs are barred very with dark gray and grayish white 
as are the limbs right out to the tips of the toes." 

Variation. — Morphometric variation among specimens 
examined is summarized in Table 2. The parietal crests in 
males may be relatively distinct and well formed (e.g., 
TNHC 31343, or they may be essentially absent (e.g., TNHC 
31342); this crest is always relatively distinct in females. 
The dorsal pattern usually consists of a contrasting array 
of dark brown or black blotches over a pale brown ground 
color, but some specimens (e.g., TNHC 46261 ) have a nearly 
uniform, dull brown dorsimi. Nearly all specimens of 



this species bear a distinctive, highly contrasting marbled 
pattern on the venter; this pattern is present, but less 
distinct in a few specimens (e.g., TNHC 46261, USNM 
297515, AMNH 69626). The middorsal cream stripe is 
present in all individuals, but may be incomplete (e.g., 
AMNH 69627). 

Etymology. — The name sij^iiifcr Latin, meaning bearing 
marks, refers to the ventral coloration of this species. 

Reproductive biology. — The tadpole of B. signifcr is 
unknown. Zweifel (personal communication) found calling 
males in a "weedy, shallow, muddy roadside pool" cin 25 
June 1962; also found in this pool were ElnchiitocU'is sp. 
and Lcptodncti/liib in^ulnniiii. 

Distribution and ecology. — This species is known 
from each of the disjunct areas of Tropical Dry Forest 
(Campbell and Lamar, 2004:47, color map 6) along the 
Pacific Coast of Panama (Fig. 13). These regions are located 
in the vicinity of city of David, and the Province of Veraguas 
eastward toward the Canal Zone. 



MORPHOMETRIC ANALYSES 

Overall morphometric variation fc)r all species is 
summarized in Table 2. Two Principal Components 
Analyses were conducted: one with log-transformed data, 
and one using residuals of each variable regressed on SVL. 
These analyses produced similar results (not presented 
here) wherein the first PC had the greatest eigenvalue 
(and accounted for the majority of the variation explained ) 
and represented an overall size axis. Despite reasonably 
high loadings for variables such as TYMP and PARW on 
the second PC, plots individual scores for each specimen 
did not clearly distinguish among the species. As could be 
expected, Biifo coccifer showed the most overall variation, 
while the smallest species (6. pisiiniiifi) was somewhat 
distinct from the largest species (B. ihnrrai) along the 
size axis (PCI). Considered together, the results of these 
analyses indicate that, with the exception of overall size, 
there has been relatively little morphometric differentiation 
among the species in this group. 

Variable selection procedures (forward, backward, and 
stepwise) for LDA suggested retention of all nine variables. 



i.e., residuals of eight variables regressed on SVL and 
log-transformed SVL. The cross-validated classification- 
matrix from the LDA is shown in Table 3. The majority 
of individuals of all species were correctly classified. 
Individuals of Biifo sigiiifer and B. porteri were misclassified 
inconsistenth' among all other species. Biifo pisimiiis and 
B. cycladcn had the greatest percentage (86%, 80'v,) of 
correctly classified individuals, respectively. The small 
size of 6. pisiiuuifi likely is responsible for this high degree 
of correct classification, but we note the 11 individuals of 
the quite variable species B. coccifer were misclassified as 
B. pifiinius; this result suggests that small B. coccifer are 
morphometrically similar tti B. pisiiniiis. Similarly, the 
relatively large size of B. ibarrai likely is responsible for 
the relatively high percentage (79' ,',) of correctly classified 
individuals of that species (and the very few individuals of 
other species that were identified as B. ibarrai). In general, 
these results are consistent with the results of the PCA. 
Despite its wide range of overall variation (Table 2), 77% 
of indi\iduals of B. coccifer were classified correctly. 



Table 3. Cross-validated classification-matrix and overall correct classification percentages (rounded to nearest integer) from Linear 
Di.scriminant Analysis based on residuals of eight morphometric variables regressed on SVL and log transformed SVL from all species in the Bufo 
coccifer Group; analysis includes only adult males. Values in boldface indicate number of individuals correctly classified. 























Per 


cent correct 




N 


B. coccifer 


B. cycladcii 


B. ibarrai 


B. pi: 


isiiuif 


B. 


sif;nifcr 


B. porteri 


cla 


ssification 


B. coccifer 


106 


82 


4 


2 




11 




4 


3 




77 


B. cyctaden 


30 


1 


24 







2 




3 







80 


B. ibarrai 


79 





2 


62 









2 


3 




79 


B. pi'^aiiiu^ 


21 


1 


1 







18 




1 







86 


B. aignifcr 


21 





2 


1 




1 




16 


1 




76 


B. porleri 


43 


3 


1 


1 




5 




1 


34 




79 



Toads of the Bufo coccifer Complex 



23 



Table 4. Voucher numbers and locality int'omiation for specimens o( Bii/o used in the molecular 
analyses. GenBank accession numbers are given for I6S and cyt-b. respectively. 



Taxon 



Voucher No 



Local itv 



GenBank No. 



B. ihcinai I 
B. iharrai 2 
B. coccifer HS 
B. coccifer Nl 
B. coccifer HO 
B. coccifer CR 
B. cycliulen 
B. conifenis 
B. valHceps 
B. mcirimis 



UT.^-ENS 10270 
UTA-.IAC 19612 
KU 290030 
SDSNH-ARH0I3 
USNM 547980 
TC'WC 8399S 
UTA-JRM 4607 
MVZ 203775 
MZFC-JRM 3868 
UTA A-50864 



Honduras: Ocotepeque 
(iuatemala: Quiche 
HI Salvador: Morazan 
Nicaragua: Ometepe Is. 
Honduras: Gracias A Dios 
Costa Rica: San Jose 
Mexico: Guerrero 
Costa Rica: Cartago 
Mexico; Veracruz 
Guatemala: Zacapa 



.A'i'927854,.'\Y927S6l 
.'\Y927S55, AY927862 
.•\Y927S56, AY927S63 
.•\Y927857. A>'927864 
.•\'i'92930I.AY929303 
.^929302. AY929304 
AY927858. AY927865 
U52800. AY008255 
AYO0821I.AYOO82I2 
AY927860. AY927867 



MOLECULAR ANALYSES 



Sequences of 547 and 410 base pairs were obtained 
for 16S and cyt-b mtDNA genes, respectively; specimen 
information and GenBank Accession numbers are listed 
in Table 4. These sections correspond to the 4004—1556 and 
16422-16818 positions for 16S and cyt b, respectively, on the 
mtDNA genome of Xeuopus [Pipidae] (Roe et al., 1985). Of 
these 956 base pairs, 765 were constant, 191 were variable, 
and 85 were considered parsimony-informative characters. 
The partition-homogeneity test results indicated that trees 
from the separate genes were not significantly different 
from one another (P = 1.0). Parsimony analysis of the 
16S region produced four trees. Parsimony analysis of 
the cyt-b region produced one tree that was identical the 
combined analysis shown in Figure 14. The differences 
among the 16S trees were the monophyly/paraphyly of 
Bufo iharrai, and the relationships among B. coccifer (sensu 
stricto) samples. Parsimony analysis of all nucleotide data 
combined evaluated a total of 2,027,025 trees, with the best 
tree score of 278 steps (Fig. 1 5) and the worst tree score of 
373 steps. The frequency distribution of trees scores had 
a mean of 351.60 steps (sd = 14.8; g, = 1.20; g, = 0.94). The 
hLRTs selected the TrN + I + G model (Tamura and Nei, 
1993) as the most significant (p < 0.01; -InL = - 2612.9934) 
with base frequencies of A = 0.3071, C = 0.2448, G = 0.1620, 
T = 0.2860. The substitution rate matrix was: A-G = 6.4270, 
C-T = 12.1788, all others were equal to 1; the proportion 



6. valliceps 
B. marinus 
B. coniferui 

B. cycladen 

B. ibarrai 
Guatemala 

5. ibarrai 
Honduras 

B. coccifer 
Nicaragua 

6. coccifer 
Honduras 

S. coccifer 
El Salvador 

B. coccifer 
Costa Rica 



Fig. 14. Phylogenetic relationships among samples used in 
molecular analyses; see Table 4 for specimen information. Topology 
is shown from the maximum-parsimony exhaustive search. Bootstrap 
values greater than 50' ». (MP /ML) are shown for each analysis above 
branches and decav indices are shown below. 





98/99 












100/82 










91 


89 






10 












11 






2 
97/74 












2 






61 






7 










1 








51 






1 





Table 5. Pair-vvisc sequence divergence of 1 6S and cyt b combmed for specimens ofBii/o used in the molecular analyses. Absolule distances are shown above 
the diagonal, and Tamura-Nei corrected distances are below the diagonal. 



Taxon 


1 


"> 


3 


4 


5 


6 


7 




8 


9 


10 


1 . B- il'<urrLtt 1 




6 


19 


17 


18 


19 


35 




70 


107 


113 


2. B. iharrai 2 


0.0064 


— 


17 


15 


16 


19 


34 




74 


105 


113 


3. B. coccifer ES 


0.0204 


0.0182 


— 


4 


3 


8 


40 




72 


106 


115 


4.B. coccifer m 


0.0184 


0.0162 


0.0043 


— 


3 


10 


40 




72 


101 


112 


5. B. coccifer HO 


0.0195 


0.0173 


0,0032 


0.0032 


— 


9 


40 




74 


107 


116 


6. S coccifer CK 


0.0205 


0.0205 


00085 


0011)7 


0.0097 




44 




75 


108 


117 


7. B cycladen 


0.0832 


0.(070 


0.0440 


0,0445 


0,0444 


0,0486 






83 


108 


114 


8. B. conifenis 


0.0791 


0.0841 


0.0816 


0,0824 


0,0848 


0,0855 


0O950 




— 


106 


124 


9. B. valliceps 


0.1261 


0.1232 


0.1253 


0.1196 


0.1277 


0,1276 


0.1269 


0.1 


235 


— 


109 


10. B. marinus 


0.1325 


0.1325 


0.1358 


0.1332 


0.1383 


0. 1 38 1 


0. 1 343 


0.1483 


0.1292 


— 



24 



SciENTii K" Pai'i:ks, Natural Hisioky Mushum,Thi-: University oi- Kansas 



of invariable sites was I = 0.5899 and the gamma shape 
parameter was G = 0.4906. A ML heuristic search produced 
a tree with a score of In L = -2609.8015 that was slightly 
different from the MP tree. In the ML tree (not shown), the 
B. ibarrai samples were basal to the remaining samples of B. 
coccifer (sensu lato); however the nodes in the ML analysis 
differing from the MP analysis were not supported in 
the ML bootstrap analysis. The ML bootstrap values are 
shown for nodes supported by more than 50% in Figure 
14. Sequence divergences for the combined 16S and cyt b 
sequences are shown for each sample in Table 5. 

The phylogenetic hypothesis generated by our 
data (Fig. 14) supports recognition of a monophyletic B. 

LITERATURE CITED 



coccifer Group that contains both B. coccifer and B. ibarrai 
(contra Blair, 1972). Our data indicate that both B. ci/cladcii 
and B. ibarrai render B. coccifer (sensu lato) paraphyletic. 
This topology, and the morphological distinctness of both 
B. ci/cladeu and B. ibarrai (Mendelson, 2001; this paper), 
suggest that B. coccifer (sensu lato) represents more than one 
species. Our proposed taxonomy recognizes six species in 
this group, one of which (B. C}/cladc}i) was here recovered 
as the sister to (B. coccifer + B. ibarrai). Knowledge of the 
phylogenetic positions of B. porteri, B. pi^iuiiu^, and B. 
^i^'^iiifer must await the collection of additional samples 
and molecular data from these species. 



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26 



SciENTiiic Papkrs. Natlral Histor'> ML'siaM.Tiiii Universh^ oi Kansas 



APPENDIX 
Specimens examined 



Note that data presented here has been taken verbatim fnmi 
museum catalogs and tags. We have not attempted to correct various 
spellings, nor to alter original estimates of distance and elevation. We 
do not consider it reasonable to re-estimate primarv locality data. 

Biifo coccifer: Costa Rica: Ai .muei a: Rio CIrande (LACM 594.'$2). 
PuNTARtNAS: 1.5 km W Barranca, 20 m (KU fi.S.'iSl); 4 km WNW Esparta, 
45 m (KU 65397-90, 65393, 65397-98, 65400, 6.5404-05, 65414). San 
Jost: Guadalupe, 1190 m (KU 65379); Esca/ii, 1100 m (KU 65380). 
Guatemala: Cimqiumii a: above headijuarters of Finca San Jose, ca. 6.0 
km SE Concepcion l.as Minas (UTA A-38119); Finca San Jose, ca. 6.0 
km SE Concepcion Las Minas on rd to Las Presas (UTA A-381 23-26); 
Concepcion Las Minas, Valle Arriba (UTA A-47566); 2.6 mi N intersection 
of CA-12 and CA-10 (UTA A-38120-21); Jalaca: Volcan Jumay (UTA 
A-47591). RtTALHULEU: 3.2 km N Champerico (UTA A-25814-'23); 3.7 
km N Champerico (UTA A-29021-25). Santa Rosa: El Oratorio, 3.0 
km E CA-8 (UTA A-38143). Hi Salvador: Chalatengo: 16.5 km WNW 
Chalatengo (KU 184659-62); 10 km NE La Palma, Canton Las Pilas 
(KU 184663-69), Cuscatlan: Tcnancingo, Rio Quezalapa (KU 184572 
i73); 7 km W Cojutepeque, 2900 ft (KU 97495); 8 km W Cojutepeque, 
2850 ft (KU 97499-504); Cojutepeque, 3220 ft (KU 97505-09); 0.5 km SE 
Cojutepeque, 2520 ft (KU 97510-11); 2 km E Cojutepeque, 3250 ft (KU 
97496). l.A Libtrtad: 10 mi NW Santa Tecla (KU 41411-29); 5 mi W Colon 
(KU 97473-86); 5.4 mi W La Libertad, 680 ft (KU 97487-92); San Bartolo, 
11 km E San Salvador, 2595 ft (KU 97493). La Union: Laguna Olomega 
(KU 184574). M(1ra/an: 12 km NE Perquin, Canton, El Zancudo (KU 
184578-609); grounds of Hotel Perquin Lenca, Perquin, 1150 m (KU 
290030). San Salvador: 1 mi NW San Salvador (KU 42613-24); Instituto 
Tropical (KU 61871-74); 16 km E San Salvador, 2880 ft (KU 97494); 3 km SE 
llopango. Canton A.sino (KU 184575-77, 184670-74); San Salvador, Cuidad 
Universitaria (KU 184612-58, 184715-16). Santa Ana: Rancho San Jose, 
800 m (KU 65372). San Vicenit;: 1 km E El Carmen (12 km E Cojutepeque), 
2620 ft (KU 97497); 6.5 km E El Carmen (17.5 km E Cojutepeque), 2700 
ft (KU 97498). Honduras: Ciiolltra: 6.8 mi S Prespire, 380 ft (TNHC 
31461); Choluteca (USNM 167195); 1.5 km NW Choluteca, 170 m (KU 
65375-76); San Francisco de Colon, 1 130 m (KU 65374). El Paraiso: 4.4 
mi SW Santa Maria, 1960 ft (KU 97512-13). Francisco Mor,-\/an: 10.3 mi 
S La Vente, 530 ft (TNHC 31454); vie. of Tegucigalpa (LSUMZ 24133-34). 
Ckacias A Dios: Rus Rus, 60 m (USNM 547977, 547980); Mocoron (UTA-CJF 
1883-84). Olancho: 1 km NW Catacamas (LACM 47973-74, 21584-89); 
Catacamas, 460 m (KU 194214); ca. Dulce Nombre de Culmi, 600 m (KU 
194213). Valle: 1 mi E Goascoran (L.A.CM 47975-77); 3 mi E Goascoran 
(LACM 47978-82); 5.2 mi SE Jicaro Galan, 250 ft (TNHC 31456, 31459). 
Mtxico: Chiapas: road from Tapachula to Puerto Madero. Oaxaca: O.I mi 
S jet 185 & 190, on 185 (LACM 38160-61 ); Juchitan (USNM 51 175); 4 mi S 
Juchitcin, 120 ft (KU 974.34-42,); 1.5 mi N Juchitan, 120 ft (KU 97443); 3 mi 
N Juchitan, 120 ft (KU 97444-17); 0.5 mi S Juchitan, 140 tt (KU 97448-50); 5 
mi N Juchitan, 110 ft (KU 97451-62); Juchitan, 9 mi 1-; jet hvvys 190 and 1S5 
(TNHC 30968); Hwy 18,5, 14.2 mi N jet Hvvy 190 ( INHC 5,3682, 53684);) mi 
NW Zanatepec (TNI IC 31 .338); 20 mi W Zanatepec (TNI IC 27161 ); 3 mi W 
Zanatepec (TNHC 271 65); 10 mi W Zanatepec (TNI IC 27292); Zanatepec 
(TNHC 27300); 17 mi E Tehuantepec (TNHC 27282, 27284, 27286, 27288, 
27290, 27305); 40 mi E Juchitan (TNHC 27291 ); edge of Tepanfepec (TNHC 
2729,3,27295,29297), Nicarac;ua: Granada: shoreofl.agode Nicaragua, 
ca 2 mi from Granada (LACM 67585). Man.-u.la: Managua, S shores 
of Lake Managua (LACM 28165-69); Los Kobles (LACM 37957-,5S, KU 
17.3951-53, 173955-.56); 2-5 km STipitapa(KU 173958), Rivas: Nandaime, 
400 ft (KU 97547). Zi i aya Noktl: Leicus Creek at La Tronquera (NIPCX) 
Lumber Plant), .56 mi NW Puerto Cabeza (l,At:M 14580S); Slilma Sia, 16 
km SE Waspam, Comarca de El Cabo (LACM 145813). 

Bufo cyclailen: Mfxico: Ck trrtro: near Palo Blanco (FMNH 99682, 
99686, UIMNT I 248,34-,38); Xaltianguis (UIMNH 24833); Agua del Obispo, 
KM 350-351 (UlMNl I 248,39); Agua del Obispo, KM 357 (UIMNH 24840); 



4 km bevond Agua del Obispo (UIMNH 24841^5); near Agua del Obispo 
(FMNH 99684-85, 99687-90, 105394, 107984-91, USNM 11548283); 
Agua del Obispo (UIMNH 24846, 24848-50, 24875-76, 57143); Agua del 
Obispo, 980 m (KU 86672-73); Agua del Obispo, 2900 ft (UMMZ 115357 
1 6 specimens]), Oax.aca: 3 mi S Putla (UIMNH 57144-51), 

Bufo ibnrrni: Guatemala: Baia Vi hai'az: ChiIasco(UTAA-47567-69); 
circa 5 km S Chilasco, 1800 m (MVZ 143379); 8 km ESE Chilasco, Finca 
Miranda, 6500 ft |198] m| (MVZ 150931); 50,2 km NW El Rancho (UTA 
A-=.016); CA-14, 29,0 mi |46,7 km] NW El Rancho (UTA A-.5049); CA- 
14, 50,2 km NW El Rancho (UTA A-5015): 4,8 mi (7,7 km] SSE Puruiha, 
Plantacion Santa Teresa (UTA A-74 1 7); 9, 1 mi (14,6 km] W Salama (by road 
to Puruiha) (UTA A-7432); 2,4 mi ]3,9 km] W Puruiha (UTA A-8502-<')7); 3.5 
mi ]5.6 km] W Puruiha (UTA A-30495 larvae); 3.2 km WNW Puruiha (UTA 
A-17117-18); 3.5 km W western Puruiha turnoff (UTA A- 17242- 1 7244); 
2.7 km W western Puruiha turnoff (UTA A-I7245); 3.8 km W Puruiha, 
1536 m (KU 186288-303); 7,7 km SE Puruiha, 1615 m (KU 186304); 3,8 
km W Puruiha, 1524 in (KU 190067); 4.2 km W Puruiha, 1524 m (KU 
190068-70); 3,4 km W Puruiha, 1524 m (KU 190071); 2,0 km W Puruiha 
(UTA A-38145^9); Hwy CA-17 between El Rancho and Coban, km 126 
(UTA A-43977-78); 1 km S San Geronimo (UMMZ 84083), El Quich6: 
Jovabaj (KU 186305); La Primavera, between Sacapulas and Santa Cruz 
Quiche, 6600 ft ]2012 m] (UMMZ 126307), Gi .ufmai a: Amatitlan (UTA 
A-.38144); 1 1,2 km SW Guatemala City 4600 ft [1402 m] (KU 97,595-609); 

21 km SW Guatemala City, 4480 ft 11366 m] (KU 97610-19]; Guatemala 
Citv, /one 10, 4820 ft ]1469 m] (TNHC 31384, 31387, 31390, 31392, 
31395, 31399, 31401-02, 31405, 31408, 31416-20, 31422, 31426, 31430-33); 
Guatemala City, between zone 5 and zone 15, km 2,5 (UTA A-25824): E 
side Guatemala City, zone 16, 1 km N Vista Hermosa III on road to Santa 
Rosita (UTA A-25825-32); Santa Catarina Pinula, San Miguel Buena Vista, 
1700 m (UTA A-43951, UTA A-47570-74); Guatemala City zone 15, Vista 
Hermosa IIL 1510 m (UTA A-28959-60);Parque San Jorge Muxbal 1850 m 
(UTA A-32993). HuEHLErtNANt.o: Aguacatan (UMMZ 120046); 2 km NE 
Aguacatan, 1640 m (UMMZ 120047-48); 2,8 km E Aguacatan, 1600 m (KU 
^8412-13); Huehuetenango, patio of Casa Maryknoll (UMMZ 124382); 

22 km SSW Huehuetenango (KU 116959); 3 km W I luehuetenango, 6100 
tt ]1859 m] (TNHC 29452-57); at La Libertad, 1700 m (MVZ 143343-57); 
San Pedro Necta (UMMZ 130059 larvae); circa 1 km F San Pedro \ecta, 
1615 m (UMMZ 119352). Jaiara: Jalapa (TNHC 33666-72); 8.5 km NW 
Jalapa (INHC 31442); 7.5 km WSW Jalapa, on road to Miramundo (UTA 
A-,391I4 larvae); Jalapa-Miramundo rd, at km 101 (UTA A-,381 18); Falda 
Oeste Volcan Jumav (UTA A-47,565); 1,6 mi [2,5 km] NE El Mojon (UTA 
A-38127-40); 0,7 nii'l 1 . 1 km] NE EI Mojon (UTA A-38141 ); Aserradero San 
Lorenzo (circa 12 air km NNE Jalapa), 1725 m (UMMZ 108000, 106806 
] 10 specimens], 106807 ]3 specimens], Proc.reso: Finca Bucaral (UMMZ 
10(iS08, 1395 16 larvae), S.u aiii-lqui/: 3 km W Dueiias (TNHC 31378); 1,3 
mi ]4,4 km] W Finca San Rafael Urias at Dueiias (TNHC 31344, 31379-80); 
San Antonio (CAS 70826-27); Volcan Agua (CAS 70719-825), Honduras: 
Kiiiiii. a: water supply area for La Fsperanza (LACM 45247—18) 1,5 mi 
NE La Esperanza (LACM 47992-96); I mi NE La Esperanza (LACM 
47998); La Esperanza (LACM 47998-48004); 25, 7 km NW La Esperanza, 
1340 m (USNM 523689-93); 18,1 km NW La Esperanza, 1740 m (USNM 
523694-96); 8,7 km NW La Esperan/a, 1540 m (USNM 523697); Zacate 
Blanco, 2020 m (KU 194220-21 ); ca, Miguelito, 60,3 km SF Gracias (Depto. 
Lempira), 1310 m (KU 209250, 2092,53), Llmrira: Frandique (LSUMZ 
46432, 497.38); above Villa Verde, 1280 m (KU 209240), OtxiTEPEOiiL: 12,5 
mi E Nueva Ocot..peque (LACM 47983-85); 6.5 mi E Nueva Ocotepeque 
(LACM 47986-91 ); ■•14'29.48'N, 88-46.83'W" (UTA A-52960, ,53662); Belen 
Gualcho, 1470 m (KU 194208); El Chaguiton, 1870 m (KU 194209-12; 
USNM .523712-13); Fl Volcan, 1760 m (USNM 523714-18), 

Bufo /lisiunus: Mexico: Miciioaian: 9 mi on rd between Ri'o 
Marque/ <md Cuatro Caminos (KU 62237— 11 ); 1-6 mi S Cuatros Caminos 
(LACM 37092-96); 7 mi F: Apal/ing.in, 1 100 fl (UMMZ I1,=;35,5, 112794 ]6 



Toads or tiii; Bufo coccifek Complex 



27 



specimens] ); 1 mi E Apatzingan, 1100 ft (UMMZ 115354 (2 specimens] ); 
3 mi S Lombardia (UMMZ 121 578): 6.2 mi E Apatzingan, 1100 ft (UMMZ 
115353 ]15 specimens], 233723). 

Bufo porteri: Honduras: Oim.wacu.a: 10.9 mi N\V Siguatepet|iie 
(TNHC 31444); 6.9 mi N\V Siguatepeque, 3820 ft (TNHC 31462, 31466); 7.5 
mi NW Siguatepeque, 3500 ft"(KU 97527); 8.8 mi NW Siguatepeque (TNHC 
31468); Siguatepeque, 3500 ft (FMNH 4612-13); 9.8 km SW Siguatepeque, 
1700 m (USNM 523683); 9.8 km SW Siguatepeque, 1950 m (USNM 
523684 );Montana de Comayagua above Ri'o Negro, 1530 m (KU 209247). 
Francisco MokazAn: Morizan, 21.4 km SE San Antonio, 4820 ft (TNHC 
31446, 31450, 31452, 31454); 17.1 mi S Tegucigalpa, 4900 ft (TNHC 31455); 
Cerro Cantagallo, 1840 m (USNM 523686); Monte Crudo, nr EAP, 6000 ft 
(AMNH 54757); La Montanuela, above Table Grande, above EAP (AMNH 
54758-59): Uyuca, above EAP, 5800 ft (AMNH 54760): Cerro Uyuca, 1900 
m (KU 209254): Uvuca, above EAP, above Tatumbia, ca. 5300 ft (AMNH 
54761): slopes of Uvuca, 4500-5000 ft (AMNH 54822); 5.5 mi SW Valle 
de Angeles (LACM 47970-71): 4.7 mi SW San Juancito (LACM 47972); 
8.6 mi NW Comavaguela (LACM 59426-31 ); 5 km W Zambrano, 1635 m 
(KU 65373); Cerro Uyuca (LSUMZ 45433, 45439-10, 45456, 46400, 46427, 
49737): Cerro La Tigra (LSUMZ 45436, 45444, 45452): Cerro La Tigra, 1840 



m (KU 194215-19, 209249); 6 mi NE Escuela Panamericana, Cerro Uyuca, 
5400 ft (KU 97514-18); 6 mi NE Escuela Panamericana, Cerro Uyuca,' 5200 
ft (KU 97519-26); W slope Cerro Uyuca, 1750 m (KU 103220); W slope 
Cerro Uyuca, 1650 m (KU 103221); Parque Nacional La Tigra, above San 
Juancito, 2100 m (KU 192294); El Hatillo (LSUMZ 45438, 45441, 45446, 
46418; LACM 72072). La P.'\z: Marcala (LSUMZ 46396-98, 46401, 46405, 
46407, 46412-14, 46420, 46422-24, 46426, 46428, 46431, 464,34-35, 46442, 
46445, 46448-51, 46453-55); Santa Elena, 1750 m (KU 194222); Sierra de 
Montecillos, about Tutule, 1750 m (KU 209244). 

Biifo signifer: Panama: Canal ZoNt: no further data (TNHC 46261 ). 
CuiRiQLii: Cerro Colorado, Escopeta Camp, ca 23 km NNE San Felix, 900 m 
(USNM 297511-15, 297517-21); Cerro Bollo, 3.5 km E of Escopeta Camp, 
1800-1850 m (USNM 297516, 297522); 7 mi N and 2 mi W David (AMNH 
69626); 7 mi E Concepcion (AMNH 69627); Cerro Bollo, 3.5 km E Escopeta 
Camp, ca. 1800 m (USNM 297522); 2.5 mi NE David (TNHC 31340-43). 
Cocle: Agua Dulce (UMMZ 167438): El Valle, 2000 ft (ANSP 23418-19); 
El Valle de Anton (AMNH 59634, 59637); 16 km S and 9 km W Penonome, 
30m(KU 1 1 5359-61 ); 3.2 km WAguadulce, 15 m (KU 115362). Herkera: 
3 mi SW Pan-Am Hwy on rd past Potuga (UMMZ 167373); Jacinto, 2250 
ft (ANSP 22341-14); Vei^guas: 14 km NE Sona, 75 m (KU 95432). 



3 2044 072 228 646 



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ing the Exchange Librarian, The University of Kansas Libraries, Lawrence, Kansas 
66045-2800, USA. Available back issues of the University of Kansas Science Bulletin 
may be purchased from the Library Sales Section, Retrieval Services Department, 
The University of Kansas Libraries, Lawrence, Kansas 66045-2800, USA. Available 
issues of former publication series. Scientific Papers, and Special Publications of 
the Natural History Museum can be purchased from the Office of Publications, 
Natural I listory Museum, The University of Kansas, Lawrence, Kansas 66045-2454, 
USA. Purchasing information can be obtained by calling (785) 864-4450, fax (785) 
864-5335, ore-mail (kunhmO'ukans.edu). VISA and MasterCard accepted; include 
expiration date. 



SiKii s Editok: William I.. Duellnian 



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